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Received: 8 August 2017    Accepted: 17 December 2017

DOI: 10.1111/jen.12489

ORIGINAL CONTRIBUTION

Diversity and abundance of lepidopteran stem borers and their

host plants in Ethiopia

M. Goftishu1,2  | Y. Assefa2,3 | A. Niba4 | C. Fininsa1 | B. P. Le Ru5,6

1
School of Plant Sciences, Haramaya
University, Dire Dawa, Ethiopia Abstract
2
Department of Zoology and Lepidopteran stem borers are among the most important insect pests of maize, sor-
Entomology, University of Fort Hare, Alice,
ghum and sugarcane in sub-­Saharan Africa. Except for Chilo partellus, the other stem
South Africa
3 borer pests in Ethiopia are indigenous to Africa and are assumed to have coevolved
Department of Crop Production, University
of Swaziland, Luyengo Campus, Swaziland with some native grasses and sedges in the natural habitat. In addition to pest species,
4
Department of Biological and Environmental natural habitats harbour diverse non-­economic stem borer species, some of which are
Sciences, Walter Sisulu University, Mthatha,
South Africa new to science. However, with the growing threats to natural habitats, some non-­
5
IRD/CRNS UMR IRD 247 EGCE, Laboratoire economic stem borer species may switch or expand their host ranges to include culti-
Evolution Génomes Comportement et vated crops and evolve as “new” pests. Besides host switch, some of the unknown
Ecologie, Comportement et Ecologie, CNRS,
Gif sur Yvette Cedex, France species currently limited to natural habitats may disappear. We examined the diver-
6
Université Paris - Sud 11, Orsay Cedex, sity, abundance and interactions of lepidopteran stem borers and their wild host plants
France
in five different vegetation mosaics in Ethiopia. The stem borer species diversity varied
Correspondence among vegetation mosaics and host plants. Forty-­four stem borer species belonging to
Muluken Goftishu, School of Plant Sciences,
14 different genera in the families of Noctuidae, Crambidae, Pyralidae and Tortricidae
Haramaya University, Dire Dawa, Ethiopia.
Email: mulukengoftishu@yahoo.com were recorded from 34 wild host plants and through light trap. Among these families,
Noctuidae was the highest in species richness in which 31 species were identified, out
of which 15 species and two genera were new to science. This paper discusses the
ecological interpretation of host plant–stem borer species interactions, particularly in
relation to habitat disturbances.

KEYWORDS
Africa, plant–herbivore interaction, species diversity, stem borer, vegetation mosaic, wild habitat

1 |  INTRODUCTION
Lepidopteran stem borers are among the most important insect
pests of cereal crops in sub-­Saharan Africa (Overholt, Maes &
Goebel, 2001; Kfir, Overholt, Khan, & Polaszek, 2002). In Ethiopia,
economically important stem borer pests of maize, sorghum and
sugarcane are the crambid Chilo partellus (Swinhoe) and the noc-
tuids Busseola fusca Fuller and Sesamia calamistis Hampson (Assefa,
Conlong, & Mitchell, 2006a; Demissie, Admassu, Getu, & Azerefegn,
2011; Gebre-­Amlak, 1985; Mengistu, Tefera, Assefa, & Yirefu,
2009; Wale, Schulthess, Kairu, & Omwega, 2006). However, the
distribution, relative abundance and damage level of these pests
vary greatly among ecological zones and crop species. Generally,
in Ethiopia, stem borer pests estimated to incur yield loss ranging
between 20% and 50% (Demissie et al., 2011; Getu et al., 2008;
Mengistu et al., 2009; Wale et al., 2006). In addition to cultivated
plants, these pests infest grasses and sedges (Assefa, Conlong, Van
Den Berg, & Mitchell, 2010; Goftishu et al., 2016; Le Ru, Ong’amo,
Moyal, Ngala, et al., 2006; Moolman et al., 2014; Ong’amo, Le Gall,
Ndemah, & Le Ru, 2014; Polaszek & Khan, 1998). Natural habi-
tats surrounding cereal fields in which these wild hosts grow are
thus considered as reservoirs for the stem borer pests (Moolman
et al., 2014; Ndemah, Schulthess, Le Ru, & Bame, 2007; Ong’amo,
Le Ru, Calatayud, Ogol, & Silvain, 2013) and their natural enemies
(Mailafiya, Le Ru, Kairu, Calatayud, & Dupas, 2009; Moolman et al.,
2013). Therefore, understanding the exchange of stem borer pests

J Appl Entomol. 2018;142:437–449. © 2018 Blackwell Verlag GmbH |  437

wileyonlinelibrary.com/journal/jen  
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438       GOFTISHU et al.
between cultivated and wild habitats is essential for developing
sound stem borer management strategies (Moolman et al., 2013;
Ong’amo et al., 2006, 2013).
Despite the critical importance of natural habitats to stem
borer pest dynamics (Ong’amo et al., 2014), previous studies in
Ethiopia focused on the distribution, diversity (Assefa et al., 2010;
Dejen, Getu, Azerefegn, & Ayalew, 2014; Gebre-­Amlak, 1985;
Getu, Overholt, & Kairu, 2001; Tefera, 2004; Wale et al., 2006)
and management of stem borer pest species in cultivated fields
only (Belay & Foster, 2010; Belay, Schulthess, & Omwega, 2009;
Gebre-­Amlak, Roland, & Jan, 1989; Getu et al., 2008; Tilahun &
Azerefegn, 2013; Tsehaye, Getu, & Fininsa, 2007). Very few studies
have been carried out on the lepidopteran stem borer fauna of the
wild habitats in Ethiopia, and they include surveys on the ecology
(Assefa, Conlong, & Mitchell, 2006b; Le Ru, Ong’amo, Moyal, Ngala,
et al., 2006) and systematics (Laporte, 1975, 1976; Rougeot, 1984;
Rougeot, Bourgogne, & Laporte, 1991). In the ecological surveys,
14 lepidopteran stem borer species belonging to the Noctuidae
(nine species), Crambidae (one species), Pyralidae (two species) and
Tortricidae (two species) were recovered from 14 host plant species
belonging to Poaceae, Cyperaceae and Typhaceae. The stem borer
F I G U R E   1   Map showing distribution of sampled localities in different vegetation mosaics in Ethiopia [Colour figure can be viewed at
wileyonlinelibrary.com]
systematics on light trap collected specimens described a total of
26 noctuid species: five Sciomesa, five Sesamia (Laporte, 1975,
1976; Rougeot, 1984; Rougeot et al., 1991) and 16 Acrapex spe-
cies (Rougeot, 1984; Rougeot et al., 1991). However, these species
were recently recombined or synonymized to Feraxinia, Sciomesa
and Sesamia genera (Moyal et al., 2010), thus reducing the total
number of species to eight. These results generated from very
few studies, and localities suggest that the diversity of the lepi-
dopteran stem borers in Ethiopia may be underestimated. Current
knowledge of stem borer species diversity in Ethiopia indicates a
lower ­specific and generic diversity compared to other sub-­Saharan
African ­countries such as Cameroon (Ndemah et al., 2007; Ong’amo
et al., 2014), Kenya (Le Ru, Ong’amo, Moyal, Ngala, et al., 2006;
Ong’amo et al., 2013), Mozambique and South Africa (Moolman
et al., 2014).
In the natural habitat, lepidopteran stem borer larvae were pre-
dominantly found inside thick and healthy stems of host plants
growing in wetter localities (forest roads, banks of streams or rivers,
swamps, lakes) (Bowden, 1976; Le Ru, Ong’amo, Moyal, Muchugu,
et al., 2006; Le Ru, Ong’amo, Moyal, Ngala, et al., 2006; Nye, 1960).
However, conversion of natural forests, grazing lands, woodlands and
GOFTISHU et al.
wetlands to agricultural land resulting from uncontrolled population
growth is the major threat to ecosystems and biodiversity in Ethiopia
(Abunie, 2003; Ethiopian Biodiversity Institute [EBI], 2014; Teshome,
2014). The fragmentation of these natural habitats may alter the dis-
tribution and diversity of stem borers (Ong’amo et al., 2013; Polaszek
& Khan, 1998).
Through extensive surveys, this study attempts to describe the di-
versity of stem borers and wild host plants in selected vegetation mo-
saics of Ethiopia. The study also describes the ecological interactions
of lepidopteran stem borers and their associated wild host plants to
predict the potential effect of habitat fragmentation on the evolution
of stem borer species.
2 | MATERIALS AND METHODS
2.1 | Selection of localities and description of the
vegetation mosaics
Surveys were conducted in five major vegetation mosaics of
Ethiopia (Figure 1) between November 2004 and December
2004, September 2006 and October 2006 and in September
2015. The types and geographic distributions of the vegetation
mosaics were described and numbered by Friis, Demissew, and
Van Breugel (2010) and White (1983). The major vegetation
mosaics included in the study were (i) Complex of Afromontane
undifferentiated forest mosaic (Undifferentiated montane veg-
etation [No. 19a]), (ii) Evergreen and semievergreen bushland
and thicket mosaic (East African evergreen and semi-­e vergreen
bushland and thicket [No. 38]), (iii) Acacia wooded grassland of
the Rift Valley mosaic (Mosaic of edaphic grassland and commu-
nities of Acacia and broad-­leaved trees [No. 63]), (iv) Somalia-­
Masai mosaic (Somalia-­M asai Acacia-Commiphora deciduous
bushland and thicket [No. 42]) and (v) Desert and semidesert
scrubland mosaic (Semidesert grassland and shrubland [No.
54b]). Localities within the vegetation mosaics were selected
based on the presence of potential host plants and accessibil-
ity. Each locality was georeferenced (latitude, longitude and
altitude) using a GARMIN 12× portable geographic positioning
system (GPS). The localities ranged from 340 to 2,689 m above
sea level (masl), 05°55′ and 12°37′ North and 36°21′ and 41°08′
East.
2.2 | Collection and rearing of stem borers
At each locality, wild host plant habitats (i) in and around culti-
vated fields, (ii) in open patch along forest roads, (iii) on river and
stream banks, (iv) in and around lakes and (v) in swamps were in-
spected for stem borer infestation. A biased rather than random
sampling procedure was used to increase the chance of finding
borers. This is because earlier studies have reported very low
stem borer densities on wild host plants compared with those of
adjacent cultivated cereals (Gounou & Schulthess, 2004; Le Ru,
Ong’amo, Moyal, Muchugu, et al., 2006; Le Ru, Ong’amo, Moyal,
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      439
Ngala, et al., 2006; Ndemah et al., 2007). In all habitats, plant
species belonging to the Poaceae, Cyperaceae and Typhaceae
were carefully inspected for stem borer infestations or damage
symptoms such as scarified leaves, dry leaves and shoots (dead
hearts), frass or holes bored. Infested plants were cut and dis-
sected in the field. Recovered larvae were all reared until pupa-
tion on artificial diets (Onyango & Ochieng-­O dero, 1994) kept
in plastic vials (7.5 × 2.5 cm) closed with perforated plastic lids.
Recovered pupae were kept separately in plastic vials (15 × 7 cm)
closed with perforated plastic lids until adult emergence. Genus
and species of the infested plants were also identified and re-
corded simultaneously in the field. Where plant species iden-
tities could not established in the field, plant specimens were
collected, pressed and taken to herbarium of the School of
Natural Resources, Haramaya University, Ethiopia, for taxonomic
identification.
In addition to inspections of wild host plants, light traps were used
in 2015 to trap flying moths. In each locality, the ideal natural hab-
itat of stem borers was selected to position the light trap. The trap
consisted of white cloth (~3 m × 3 m) reinforced with ropes and tied
between two stands of wooden splints. The ground under the white
cloth was covered with white canvas of a similar size. Two electrical
bulbs (white light sources) were tied on a wooden splint stand in front
of the white cloth with one bulb lighting above the cloth and the other
in the middle of the cloth to attract moths. Stem borer moths gather-
ing on the white cloth and those that fall on the canvas were collected,
pinned and identified later.
2.3 | Identification of moth species and analysis of
borer species diversity
Adult stem borers were identified to species level by the fifth
author. Voucher specimens were deposited in the International
Centre of Insect Physiology and Ecology (ICIPE, Nairobi, Kenya)
Museum and in Museum national d’Histoire naturelle (MNHN,
Paris, France). Identified stem borers were grouped according to
their respective host plants and vegetation mosaics for analysis
of stem borer species diversity. Data of previously recorded stem
borer species (Laporte, 1975, 1976; Rougeot, 1984 and Rougeot
et al., 1991) were also included in an attempt to catalogue the
diversity of the stem borer species in Ethiopia. Data from each
­vegetation mosaic were used to compute diversity indices de-
scribed by Magurran (2005) (Tables 1 and 2). Species richness (S)
was one of the indices computed. This index is based on the pres-
ence, rather than the relative abundance. This conceptually easy
index is widely used in studies related to species ecology and bio-
geography. Other indices computed were the alpha (α) and Berger–
Parker dominance (d) indices. Alpha diversity index (α) captures
the mean species diversity in sampled subunits and calculated as
a weighted generalized mean, while Berger–Parker dominance (d)
index represents the proportional abundance of the most domi-
nant species relative to the abundance of all species together for
a sample unit.
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440       GOFTISHU et al.

T A B L E   1   List of stem borer species

Vegetation mosaic no.
collected from wild host plants in different
Family No. Stem borer species 19a 38 42 54b 63 vegetation mosaics in Ethiopia

Noctuidae [1] Acrapex mondogeneta 5

sp. nov.
[2] Busseola fusca 128 128 15
[3] Busseola sp. nov. 1 5
[4] Busseola sp. nov. 3 41 21 20
[5] Busseola s.l. 2 sp. nov. 74
2
[6] Busseola s.l. 2 sp. nov. 15
3
[7] Busseola segeta 77 17
[8] Feraxinia jemjemensis 324 14
[9] Feraxinia secata 2
[10] Manga nubifera 93
[11] Pirateolea piscator 6 10
[12] Pirateolea sp. nov. 3 10
[13] Sciomesa mesophaea 1
[14] Sesamia calamistis 16 45
[15] Sesamia cretica 8
[16] Sesamia nonagrioides 2 91 46 110
[17] Sesamia sp. nov. 40 10 3
[18] Sesamia sp. nov. 42 48
[19] Sesamia sp. nov. 43 100
Pyralidae [20] Eldana saccharina 126 50
[21] Phycitiinae sp. 2 5
[22] Phycitiinae sp. 16 2
[23] Phycitiinae sp. 19 35
Crambidae [24] Chilo partellus 20
[25] Crambidae sp. 4
Tortricidae [26] Tortricidae sp. 13
[27] Tortricidae sp. 12 1
[28] Tortricidae sp. 13 1 2
[29] Tortricidae sp. 14 10
[30] Tortricidae sp. 15 12 2
[31] Tortricidae sp. 16 1
[32] Tortricidae sp. 17 4
Taxa (S) 17 13 6 1 10
Individuals (n) 729 581 117 8 338

Numbers under respective columns of vegetation mosaics are the total number of individuals of each
species.

(H′2) and generality (Gqw) at the network/community level. At the

2.4 | Constructing stem borer–host plant
ecological network
Host plant–stem borer interaction networks were analysed by “bi-
partite” package (Dormann, Gruber, & Fründ, 2008) in R-­software
(R Core Team, 2016). To characterize network structures for each
vegetation mosaics, we calculated the network specialization index
species level, we computed the species specificity index (standard-
ized Kullback-­Leibler distance, d′i ). These three indices were selected
among the multitude of network indices because they describe net-
work properties quantitatively and are relatively robust against vari-
ation in sampling effort and network size (Bersier, Banasek-­Richter,
& Cattin, 2002; Bluthgen, Menzel, & Bluthgen, 2006; Kaiser-­Bunbury
GOFTISHU et al. |
      441

T A B L E   2   List of stem borer species collected with light traps in analysis because no meaningful network index could be calculated
different vegetation mosaics in Ethiopia when a trophic level has only a single species.

Vegetation mosaic no.

Family Stem borer species 19a 38 42 54b 63 3 | RESULTS

Noctuidae Acrapex sp. nov. 95 1
3.1 | Stem borer species diversity
Acrapex mondogen- 82 5
eta sp. nov. A total of 2002 stem borers were collected both from host plants
Acrapex sp. nov. 97 12 and light traps. From this collection, a total of 44 stem borer species
Acrapex robe sp. 13 were identified (Tables 1 and 2). Identified stem borers included both
nov. known and unknown species. The unknown stem borer species could
Acrapex sp. nov. 1 only be identified to family or genus levels and are presented here
101
in their corresponding lowest taxonomic units. The identified species
Acrapex soyema sp. 1 belonged to four families as follows: Crambidae, Noctuidae, Pyralidae
nov.
and Tortricidae (Tables 1 and 2). Among these families, Noctuidae
Busseola fusca 7
was the highest in species richness in which 31 species were iden-
Feraxinia 26 [6]
tified. Stem borer species identified in the Noctuidae belonged to
jemjemensis
nine genera (Acrapex, Busseola, Feraxinia, Manga, Pirateolea, Sciomesa,
Feraxinia secata [13]
Sesamia, New Genus 4 and New Genus 19). Of the nine Noctuidae gen-
Manga nubifera 2
era, Sesamia was the richest with seven species followed by Acrapex
Sciomesa [5] and Busseola each with six species. The other important genus was
apexangula
Sciomesa represented by five species followed by Feraxinia (two spe-
Sciomesa boulardi 27 [3]
cies), Pirateolea (two species) and Manga (one species; Tables 1 and
Sciomesa congitae [2] 2). Four species were identified in the family Pyralidae. There were
Sciomesa excels [2] two species in the family Crambidae and seven species in the family
Sciomesa 2 [3] [1] Tortricidae.
mesophaea
Sesamia calamistis 1 1
Sesamia [3]
3.2 | Abundance and diversity of stem borer species
nonagrioides among vegetation mosaics
Sesamia sp. nov. 41 2 Abundance and diversity of stem borer species were analysed at
New Genus 4 sp. 7 the vegetation mosaic level to obtain an estimate of the ecological
nov. 6
preference of the different species. The highest number of species
New Genus 19 sp. 1
(S) was recorded in Afromontane mosaic (24) where a total of 915
nov. 1
individuals were collected (Table 3). In the East African mosaic,18
Taxa (S) 11 7 1 0 4
species were identified from 598 individuals, followed by Acacia
Individuals (n) 186 17 1 0 25
wooded grassland of the Rift Valley mosaic where 12 species were
Numbers under respective columns of vegetation mosaics are the total identified from 363 individuals. The remaining two vegetation mosa-
number of individuals of each species. Part of the collections made by ics, Somalia–Masai and the semidesert grassland and shrubland had
Laporte (1975, 1976), Rougeot (1984) and Rougeot et al. (1991) are en-
seven and one species each from 118 and eight individuals, respec-
closed in parenthesis [].
tively (Table 3).
Stem borer species abundance varied in respective vegetation mo-
& Bluthgen, 2015). The specialization index describes the level of saics. The Afromontane mosaic (α = 4.52) was dominated by Feraxinia
specialization, or selectiveness, of an entire network (Bluthgen et al., jemjemensis (Figure 2b, species SB 11) which comprised 39% (Table 3)
2006; Kaiser-­Bunbury & Bluthgen, 2015), ranging from 0 (low selec- of the total collection in the vegetation mosaic followed by B. fusca and
tiveness) to 1 (high selectiveness). The species specificity index ex- Acrapex mondogeneta sp. nov. (Le Ru, Capdevielle-­Dulac, C., Musyoka,
presses how selective a given species is relative to available resources B. K., Pallangyo, B., Njaku, M., Goftishu, et al., 2017). The stem borer
(Bluthgen et al., 2006; Kaiser-­Bunbury & Bluthgen, 2015), ranging be- species community in the East African evergreen and semievergreen
tween 0 (perfect opportunist) and 1 (disproportionate specialist). Gqw bushland and thicket mosaic (α = 3.50) was dominated by B. fusca
is the mean effective number of plant species per herbivore species (Figure 2c, species SB 2) which constituted about 21% (Table 3) of the
(Bersier et al., 2002; Kaiser-­Bunbury & Bluthgen, 2015). Furthermore, total collection in the vegetation mosaic followed by Eldana saccharina
the semidesert grassland and shrubland mosaic (n = 1) with only a and Manga nubifera. The Acacia wooded grassland of the Rift Valley
single plant to herbivore species interaction were excluded from the mosaic (α = 2.39) was dominated by Sesamia nonagrioides (Figure 2d,
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442       GOFTISHU et al.

T A B L E   3   Number of individuals, species richness, alpha and recorded from several wild host plant species (Table 4). These
berger-­parker dominance index of the full data set (light traps and species constituted more than 34% of the total number of stem
wild hosts) borers collected from wild host plants. Among the pest species,
Vegetation mosaic no. B. fusca was the most abundant with 271 individuals collected from
Arundo donax and Pennisetum purpureum, with most of the larvae
Parameter 19a 38 42 54b 63
(91%) found on A. donax (Figure 2a, species HP 2). Busseola fusca
Number of 915 598 118 8 363 accounted 15.3% of the stem borer populations collected from wild
individuals (n)
hosts. The second most abundant pest species was S. nonagrioides
Species richness (S) 24 18 7 1 12
with 249 (Table 1) individuals obtained from six host plants from the
Alpha (α) 4.52 3.50 1.63 – 2.39 Poaceae and one Typhaceae (Table 4), although most (167; 67%) of
Berger-­Parker (d) 0.39 0.21 0.39 – 0.30 the larvae were recovered from Typha domingensis (Figure 2a, spe-
cies HP 33). It accounted for 14% of borer populations recovered
species SB 22) which constituted 30% (Table 3) of all individuals in the from wild hosts. Sesamia calamistis was collected from five species
mosaic. Similarly, the Somalia–Masai Acacia-Commiphora deciduous of Poaceae and one species of Cyperaceae (Table 4), making up
bushland and thicket mosaic (α = 1.63) was dominated by S. nonagri- 3.4% of all stem borers collected from wild hosts. Chilo partellus
oides which constituted (Figure 2e, species SB 22) 39% (Table 3) of all and S. cretica each recovered from A. donax and Sorghum halepense,
individuals in the mosaic. in that order. Individuals of these two species together makeup
<2% of the stem borer population collected from wild host plants
(Table 1).
3.3 | Abundance and distribution of stem borers
Among the non-­pest species of stem borers, F. jemjemensis was
among wild host plants
the most abundant and dominant species which accounted for more
The economically important stem borer species, namely B. fusca, than 19% (338 individuals) of the stem borer populations collected
C. partellus, S. calamistis, S. cretica and S. nonagrioides, were from wild host plants. Feraxinia jemjemensis was obtained from nine

(a)

F I G U R E   2   Stem borer–host plants

(b)
interaction networks. (a) The full data
set. (b) Complex of Afromontane
undifferentiated forest mosaic. (c)
Evergreen and semievergreen bushland and
thicket mosaic. (d) Acacia wooded grassland
of the Rift Valley mosaic. (e) Somalia–Masai
Acacia-Commiphora deciduous bushland
and thicket. In every network, lower bars
represent host plant species and upper
bars represent the stem borer species. Each
black rectangle represents the proportional
abundance of stem borers and host plants
and each grey triangle represents the
frequency of interaction between them.
Codes refer to stem borers and host plants
species names in Table S1
GOFTISHU et al.
(c)
(d)
(e)
F I G U R E 2   Continued
host plants all from the family Poaceae, with most of the larvae
found on Setaria megaphylla (37.87%; Figure 2a, species HP 29).
Eldana saccharina was also the most abundant species which com-
prised 10% (176 individuals) of the stem borer populations obtained
from wild host plants. All individuals of E. saccharina were collected
from three species of Cyperaceae, with most of the larvae (66.5%)
recovered from Cyperus papyrus (Figure 2a, species HP 9). The other
abundant non-­pest species include Busseola sp. nov. 3, B. segeta and
|
      443
M. nubifera. Busseola sp. nov. 3 was collected from five host plants
all from the family Poaceae, making up 4.60% of all populations of
stem borers recovered from wild host plants. Each species of B. seg-
eta and M. nubifera comprised 5.3% of all borer population collected
from wild host plants. Busseola segeta was obtained from three host
plants from the Poaceae only, with majority of the larvae (96.80%)
were recovered from Panicum deustum, whereas M. nubifera were
recovered from Panicum maximum only.
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444       GOFTISHU et al.

T A B L E   4   Wild host plant species from which different stem borer species collected in different vegetation mosaics in Ethiopia

Vegetation mosaic no.

Plant family Tribe Host plants 19a 38 42 54b 63

Poaceae Andropogoneae Andropogon sp. [17]

Cymbopogon citratus [5] [14]
Cymbopogon nardus [4] [8]
Hyparrhenia hirta [1]
Hyparrhenia lintonii [8] [14]
Hyparrhenia papillides [8]
Rottboellia cochinchinensis [22]
Sorghum halepense [15]
Vossia cuspidate [16]
[26]
Arundineae Arundo donax [2] [2] [11] [2]
[16]
[24]
Phragmites mauritianus [19]
Paniceae Eriochloa procera [25]
Echinochloa pyramidalis [4][11][14][16][18] [16] [16]
Panicum deustum [3] [7] [3] [7]
Panicum mapalende [19]
Panicum maximum [4] [4][7][10][21] [17]
Pennisetum clandestinum [4] [5] [8] [11]
Pennisetum macrourum [12]
Pennisetum purpureum [2] [8] [2] [4] [11] [4]
[16]
Pennisetum thonbergii [16]
Pennisetum trachyphyllum [5] [8] [9] [11]
Pennisetum unisetum [6] [14] [23]
Setaria megaphylla [8] [7]
Setaria plicatilis [8] [8]
Snowdenia polystachya [8]
Cyperaceae Cyperaceae Cyperus distans [28]
Cyperus dives [20] [20]
Cyperus fastigiatus [20]
Cyperus papyrus [20]
Cyperus rotundus [32]
Cyperus latifolius [13]
Schoenoplectus articulatus [29]
Schoenoplectus corymbosus [28] [30] [30] [14] [31]
[27]
Typhaceae Typhaceae Typha domingensis [16] [16] [16]
[19]

Values in bracket [] under respective columns of vegetation mosaics represents codes of different stem borer species as listed in Table 1 above. Check the
codes and frequency of borers.

Based on individual numbers of the collected specimens, Sesamia accounted 5.64% of all stem borer species collected from wild host
sp. nov. 43 and Busseola s.l. 2 sp. nov. 2 were also considered as abun- plants. Similarly, Busseola s.l. 2 sp. nov. 2 was obtained from three host
dant. The larvae of Sesamia sp. nov. 43 were collected from three plants from Poaceae only (Table 4) and accounted 4.17% of the stem
host plants from the family Poaceae and one Typhaceae (Table 4) and borer populations recovered from wild host plants. However, these
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      445

T A B L E   5   Network descriptors for

Vegetation mosaic no.
community level stem borer–host plant
interaction networks in selected vegetation Network descriptors All 19a 38 42 63
mosaics of Ethiopia
No. of moth species 32 17 13 6 10
No. of plant species 34 18 18 5 10
Linkage density 2.10 2.20 1.34 1.21 1.56
Shannon diversity of 3.31 2.73 2.27 1.33 2.13
interactions (H′)
Network specialization 0.85 0.83 0.98 0.98 0.97
(H′2)
Generality (Gqw) 2.40 2.79 1.41 1.32 1.64

two species were rare (cf Le Ru, Ong’amo, Moyal, Muchugu, et al.,
2006). Rare stem borer species were recorded both from light trap
(eight species) and wild host plants (18 species) in all the vegetation
mosaics.
3.4 | Diversity of host plants
The wild host plants recorded during these surveys are listed
in Table 4. Stem borer larvae were collected from 34 host plant
species belonging to three families: Poaceae (25), Cyperaceae (8)
and Typhaceae (1) (Table 4). Of 1,773 larvae collected, 76.10%
were from Poaceae, 14.37% from Cyperaceae and 9.53% from
Typhaceae.
Within the family Poaceae, high diversity of borer species was
found on Echinochloa pyramidalis, P. maximum and P. purpureum,
where each plant hosted five species of stem borers. Four species
were recorded on A. donax, P. maximum, Pennisetum clandestinum
and Pennisetum trachyphyllum. In Cyperaceae, high species diversity
(five species) was found on Schoenoplectus corymbosus. The remain-
ing plant species harboured one to three species of borers (Figure 2;
Table 4).
3.5 | Host plant–stem borer interaction network
Ecological network analysis is a valuable tool for studying species
diversity and interactions within and across trophic levels (Albrecht,
Duelli, Schmid, & Muller, 2007). Interaction networks are graphic
representations of interacting species in a community (Figure 2). A
host plant–stem borer interaction network of the complete data set
consisted of 34 plant and 32 stem borer species and are depicted
in Figure 2a (see Table S1 for species names and codes). The host
plant–stem borer interaction network descriptors of the full data set
showed high index of network specialization (H′2 = 0.85) and low index
of generality (Gqw = 2.40; Table 5), suggesting that each stem borer
species had on average less than three “effective” host plant species.
This result further revealed the majority of borer species feed on a
limited number of host plants, suggesting that these species are either
monophagous or oligophagous.
Among the vegetation mosaics, network interactions were rela-
tively diverse in the Afromontane vegetation mosaic, followed by East
African and Acacia wooded grassland mosaics as shown by Shannon di-
versity indices (Table 5). Linkage density ranged from 1.21 in Somalia–
Masai to 2.20 in Afromontane vegetation. Likewise, generality ranged
from 1.32 in Somalia–Masai to 2.79 in Afromontane vegetation mo-
saic (Table 5). High index of network specialization was observed in
all vegetation mosaic (Afromontane: H′2 = 0.83; East African: H′2 = 0.98;
Acacia wooded grassland: H′2 = 0.97; Somalia–Masai: H′2 = 0.98), show-
ing that these host plant–stem borer interaction networks at the
community level tended towards complete selectivity. Similarly, the
species-­level (d′i ) specificity of stem borers ranged from 0.20 to 1.00
(median 0.82) in the Afromontane vegetation mosaic, from 0.30 to
1.00 (median 0.72) in the East African vegetation mosaic, from 0.60 to
1.00 (median 0.96) in the Acacia wooded grassland and from 0.25 to
0.98 (median 0.94) in the Somalia–Masai vegetation mosaic (Table S2).
3.6 | Contribution of light trap collection to
species diversity
A total of 20 stem borer species all from the family Noctuidae were
collected from light traps. Of these species, 12 species were exclu-
sively collected from light traps (Table 2). Some of these stem borer
species belonging to the Acrapex genus (five species), Sesamia genus
(one species) and two genera new to science. Eight species were
found both in light traps as well as in host plants (Tables 1 and 2).
4 | DISCUSSION
Although several taxonomic studies have been carried out in the
past on stem borers (Laporte, 1975, 1976; Le Ru et al., 2014; Moyal
et al., 2010; Rougeot, 1984; Rougeot et al., 1991), many new stem
borer species were collected during this study. Of the 44 stem borer
species recorded, only 16 were identified to species level. The re-
maining stem borers could only be identified to family or genus
level; all noctuid borers (17 species) identified to the genus level
are “new” to science. Even though, this study was limited in time
and space and probably underestimated the diversity of noctuid
stem borers, the result confirms that stem borer species diversity
in Ethiopia is much higher than earlier reported. Many more species
may be reported if this kind of study covers Combretum-terminalia
 |
446       GOFTISHU et al.
woodland ecosystems of western and north-­western Ethiopia, par-
ticularly known of diverse species of grasses and sedges (Ethiopian
Biodiversity Institute, 2014).
The undifferentiated montane vegetation that mainly covers the
highlands of Central, South-­Eastern and South-­Western Ethiopia was
the richest in stem borer species diversity. The Ethiopian highlands
cover much of the countries land area (Taylor, 2015). These highlands
are source of water to rivers, streams and wetlands, and often re-
ferred as the “water tower of Northeast Africa” (Williams, Pol, Spawls,
Shimelis, & Kelbessa, 2005; Wondefrash, 2003). In wild habitats, lep-
idopteran stem borers are mainly found in thick grasses and sedges
growing in wetter localities (Le Ru, Ong’amo, Moyal, Ngala, et al.,
2006). Thus, the relative abundance of such habitats in high-­altitude
areas of Ethiopia might have contributed to the diversity of stem bor-
ers recorded. Similar to our findings, high diversity of noctuid stem
borers was reported in the high-­altitude areas of East (Le Ru, Ong’amo,
Moyal, Muchugu, et al., 2006; Ong’amo et al., 2013) and Central Africa
(Ong’amo et al., 2014). In addition to species richness, the undifferen-
tiated montane vegetation harboured many of the new noctuid stem
borer species. Majority of the newly recorded species were, however,
recovered from one locality, suggesting that such species are rare in
the region. The occurrence of rare species in the natural habitat was
frequently reported for herbivorous insects in tropical regions (Le Ru,
Ong’amo, Moyal, Muchugu, et al., 2006; Novotny & Basset, 2000;
Ong’amo et al., 2014). The presence of rare species may not necessar-
ily indicate sampling artifacts, as in the face of very intensive sampling,
the incidence of rare species are also common (Le Ru et al., 2014;
Moyal et al., 2010). The newly recorded noctuid stem borer species in
our study could possibly be endemic to Ethiopia. Similar studies con-
ducted in sub-­Saharan Africa also confirmed the presence of a high rate
of endemism in noctuid stem borers particularly in the genus Acrapex
(Le Ru, Capdevielle-­Dulac, C., Musyoka, B. K., Goftishu, et al., 2017;
Le Ru, Capdevielle-­Dulac, C., Musyoka, B. K., Pallangyo, B., Njaku, M.,
Goftishu, et al., 2017; Le Ru, Capdevielle-­Dulac, C., Musyoka, B. K.,
Pallangyo, B., Njaku, M., Kandonda, et al., 2017; Le Ru et al., 2014). As
part of the Eastern Afromontane hotspot, the Ethiopian highlands are
known for their high concentrations of endemic species (CEPF, 2012;
Mittermeier et al., 2004; Williams et al., 2005).
This study recorded 34 wild host plant species which is more
than double of that of the previous reports. Species diversity of stem
borers correlates positively with increased plant diversity (Moolman
et al., 2014). However, the majority of stem borer species had a lim-
ited host plant range. This is further supported by the result of host
plant–stem borer interaction networks that exhibited a high degree
of network specialization (H′2) and host plant specificity (d′i ) of stem
borers at the community and species level, respectively. Similar
findings were reported for other plant–insect herbivore networks
(Araujo, Grandez-­Rios, Bergamini, & Kollar, 2017; Cagnolo, Salvo,
& Valladares, 2011). From an ecological network perspective, this
means that the removal of plant species causes few changes in the
network structure, as each host plant species is connected to one or
a few species of stem borers. According to Araujo et al. (2017), this
result also means that the loss of a plant species will, in most cases,
most likely lead to extinction of the associated stem borer species
in the network. If a species is lost from an ecosystem, it is not only
the species itself that is lost, but its interactions, and the ecological
functions that result from these interactions, for example, natural
enemies (Kankonda, Akaibe, Ong’amo, & Le Ru, 2017). Even if the
network metrics revealed specialization, among vegetation mosaics,
indices of linkage density and Gqw were comparatively higher in the
undifferentiated montane than the remaining vegetation mosaics,
where relatively more intense human-­mediated habitat disturbances
occurred (EBI, 2014). High index of Gqw implies higher community
stability (Kaiser-­Bunbury & Bluthgen, 2015), whereas low Gqw could
be a result of the loss of lower trophic levels, typical characteristics of
fragmented habitats (Valladares, Cagnolo, & Salvo, 2012).
Insect herbivores switch more easily between closely related
plants (Brandle & Brandl, 2001) and typically perform better on
agricultural plants than on wild ancestors due to reductions in
secondary compounds during crop domestication (Chen, 2016).
Resource concentration hypothesis (Root, 1973) also suggests that
if plant species are more abundant or nearly pure stands, insects
are able to specialize on them and often do so. The colonization of
E. saccharina in South Africa and Zimbabwe from wild sedges onto
sugarcane confirmed this hypothesis (Conlong, 2001; Mazodze &
Conlong, 2003). All indigenous African lepidopteran stem borer
pests are assumed to have coevolved with some native grasses
and sedges and switch or expanded their host range to cultivated
plants upon exposure to anthropogenic changes (Nye, 1960;
Polaszek & Khan, 1998; Leslie, 2004). In a recent survey conducted
in Tendaho sugarcane estate, northeastern Ethiopia, that started
sugarcane production a few years before by clearing the natural
vegetation, S. cretica was found as the only economically important
sugarcane stem borer pest (Goftishu et al., 2016). Sesamia cretica
is a major pest of sorghum and sugarcane in East and North Africa
(Kfir et al., 2002; Leslie, 2004; Ezzeldin, Sallam, Helal, & Fouad,
2009). A similar survey carried out in three sugarcane estates
located in the central Rift Valley of Ethiopia also found high in-
festation of S. nonagrioides and recorded the species for the first
time as one of the major sugarcane stem borer pest in the estates
(M. Goftishu, Y. Assefa, C. Fininsa, A. Niba, C. Nyamukondiwa, C.
Capdevielle-Dulac, & B. P. Le Ru, unpublished results). Host range
expansion of S. cretica to sugarcane and the recent record of
S. nonagrioides to the pest community provide evidence of a sig-
nificant risk of emergence of other “new” pests. In our study, few
stem borer species (e.g., B. segeta and E. saccharina) currently re-
stricted to natural habitats were dominantly recorded on wild host
plants. Busseola segeta was recently reported as a potential pest of
maize in high-­altitude areas of Kenya (Calatayud, Okuku, Musyoka,
Khadioli, & Ong’amo, 2014; Ong’amo et al., 2013). According to
Calatayud et al. (2014), in some year, the infestation percentage of
B. segeta was much higher than the major stem borer pest, B. fusca.
Similarly, E. saccharina is a serious pest of sugarcane in southern
African countries (Conlong, 2001; Mazodze & Conlong, 2003).
Even if they are not yet recorded from cultivated plants in Ethiopia,
with the loss of natural habitats these species together with other
GOFTISHU et al.
stem borers that are currently limited to wild hosts may have the
potential to shift to cultivated plants.
In addition to wild stem borers, pest species of stem borers par-
ticularly B. fusca and S. nonagrioides were commonly observed among
wild host plants. A similar result was obtained in Eritrea where B. fusca
was found very common on wild hosts (Le Ru, Ong’amo, Moyal, Ngala,
et al., 2006). Contrary to our findings, earlier studies undertaken in
Central, East and Southern Africa indicated that economically im-
portant borer species were less abundant in wild habitats compared
with non-­pest species (Moolman et al., 2014; Ndemah et al., 2007;
Ong’amo et al., 2013). Wild habitats were reported as reservoirs for
stem borer pests during non-­crop seasons (Bowden, 1976; Polaszek
& Khan, 1998). However, the abundance of pest species on wild host
plants could not be used as the only decisive criterion to measure the
contribution of natural habitats as refuges for stem borer pests. To ef-
fectively determine whether wild habitats serve as real reservoirs of
stem borer pests during non-­cropping period and subsequently infest
cultivated plants during cropping season, in addition to abundance,
other factors such as dispersal ability, diapause, exchange and genetic
structure of pest species between natural and cultivated habitats need
to be quantified.
ACKNOWLE DG E MEN TS
We express our appreciation to Haramaya University and the South
African National Research Foundation (NRF) for providing the re-
search fund. We acknowledge the University of Fort Hare, Faculty
of Science and Agriculture, Department of Zoology and Entomology
for providing a Ph.D. fellowship to the first author. We are very
grateful to Professor Nigussie Dechassa, Academic Vice President
of Haramaya University (the then Research Vice President) for his
enthusiastic assistance during the study period. We also appreciate
Mr. Zelalem Gebeyehu for his help with preparing the sampling site
map.
AUT HOR CONTRI B UTI O N
MG collected stem bore specimens from the field, analysed the data
and contributed to writing the paper. BPLR collected specimens from
the field, morphologically identified adult stem borers to species level
and contributed to writing the paper. YA contributed additional speci-
mens and involved in writing the paper. AN and CF contributed to
writing the paper. All authors read and approved the final manuscript.
O RCI D
M. Goftishu  http://orcid.org/0000-0002-7194-9470
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S U P P O RT I NG I NFO R M AT I O N
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How to cite this article: Goftishu M, Assefa Y, Niba A, Fininsa
C, Le Ru BP. Diversity and abundance of lepidopteran stem
borers and their host plants in Ethiopia. J Appl Entomol.
2018;142:437–449. https://doi.org/10.1111/jen.12489