Higher diptera associated with the marsh spike-rush, Eleocharis palustris (cyperaceae), in northeastern Ohio

2007·3.9 MB·

by  B A Foote

PROC. ENTOMOL. SOC. WASH.
109(1), 2007, pp. 9-16
HIGHER DIPTERA ASSOCIATED WITH THE MARSH SPIKE-RUSH,
ELEOCHARIS PALUSTRIS (CYPERACEAE), IN NORTHEASTERN OHIO
B. A. FooTE
Department of Biological Sciences, Kent State University, Kent, Ohio 44242
(e-mail: [email protected])
—
Abstract. Information is given on 28 ofthe 50 species ofhigher Diptera that were
found in stands ofthe marsh spike-rush, Eleocharispalustris (L.) Roemer & Schultes,
in northeastern Ohio. Seasonal distributions, abundances, and larval feeding habits
of selected species are presented.
Key Words: Eleocharis, Diptera, freshwater marshes, Ohio
This is the ninth paper in a series that long and narrow (100 X 5 m) and
focuses on the species of higher Diptera consisted of a near monoculture of E.
(Schizophora) that occur in freshwater palustris, although a few individuals of
marshes in northeastern Ohio (Todd and rice cut grass, Leersia oryzoides (L.)
Foote 1987a, b; Rogers et al. 1991; Swartz, were scattered throughout the
Wearsch and Foote 1994; Larson and stand. The stand was bordered on either
Foote 1997; Keiper et al. 1998; Foote side by an undetermined species of grass
2004a, b). The present study gives survey and by narrow-leaved cattail (Typha
data obtained during one field season of angustifolia L.) at its northern end. The
collecting higher Diptera from a near- study site was mowed periodically by
monoculture stand of the marsh spike- county maintenance crews throughout
rush, Eleocharispalustris (L.) Roemer & the summer months. Water, at a depth of
Schultes, growing in a roadside drainage some 10 cm, was present during the
ditch near Kent in northeastern Ohio. spring months. Water depths receded
Information is given on seasonal occur- steadily as summer advanced, although
rence and larval feeding habits of 28 of summer rains caused temporary fluctua-
the 50 associated species. tions. By late August, no standing water
was present.
Materials and Methods
Specimens ofDiptera were obtained by
Eleocharis palustris is a widespread, sweep samples consisting of 15 back and
narrow-leaved, emergent, perennial spe- forth movements of a 15 inch diameter
cies of Cyperaceae having horizontal, standard aerial insect net through the
mat-forming rhizomes (Ball et al. 2002). stand. Sampling took place weekly for
The study stand was located in a shallow 22 weeks between 6 May and 30 Septem-
drainage ditch bordering Hodgeman ber, 2004. Nearby stands of rice cutgrass
Lane, a small road leading to the Kent and other undetermined grass species
Water Treatment Plant on the east side were swept for comparative purposes.
of the city in Portage County (41° 08' Rearings were initiated from larvae or
33"N X 81° 19' 19"W). The stand was adults collected from the host plant.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
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Rearing containers were small (5.5 X third instar being reached the following
7.0 cm) plastic jars from which the late April or May. Mature larvae mi-
bottom had been removed. The jar was grated to the rhizomes where they
then inserted into a Petri dish containing fonned puparia in May. The pupal
a layer of moist peat moss. The top of period ranged from 10-15 days under
the jar was covered with fine-mesh laboratory conditions. Stem infestation
nylon. A small pellet ofmoistened honey rates were quite low, with 5 samples of30
and yeast was affixed to the wall of the field-collected stems each having emer-
jar to serve as food for the adults. gence rates ranging from to 5%.
Once larvae were obtained, they were Infested stems rarely produced inflores-
transferred to Petri dishes containing cences.
—
fragments of the host plant or a field- Diplotoxa inclinata Becker. This
collected sample of detritus or algae. strictly Nearctic species has a transconti-
Larvae were killed in hot water and nental distribution in North America
preserved in 70% ethanol. Puparia that and was abundant in the study stand
produced adults were placed in small between early May and early September.
plastic vials and pinned beneath the It was usually the first species of
emerged adults. Voucher specimens are Diplotoxa to appear at the study site.
deposited in the insect collection of the Based on evidence obtained from the
author. sweep samples and from laboratory
rearings, it was an univoltine species.
Results
Mated females confined with Eleocharis
A total of 875 specimens from 35 in breedingjars readily deposited eggs on
genera and 50 species were collected the stems, although a few eggs were also
from the stand of Eleocharis during the deposited on the underlying peat moss.
summer of2004 (Table 1). In the follow- Most ofthe eggs were placed at the base
ing annotated list, 28 species are covered ofthe stemjust above the peat moss, but
in greater detail with respect to their a few were attached at greater heights.
seasonal occurrence, relative abundance, None was placed on inflorescence. The
and larval feeding habits. incubation period lasted 4-5 days. New-
ly hatched larvae crawled down the stem
Annotated List of Selected Species and entered young shoots at the base of
the plant where they began feeding on
Family Chloropidae
— the stem tissue. Older larvae moved
Chlorops obscuricornis Loew. This downward within the stem and penetrat-
Nearctic species is a stem borer of ed into the rhizomes attached to the
Eleocharis, having been reared from crown of the plant. They then continued
stems of E. palustris and E. ohtusa to feed within the meristematic tissue of
(Willd.) (Wearsch and Foote 1994). It the rhizome. Overwintering occurred as
was an univoltine species having a flight inactive second instars, and feeding re-
period from late May to early August. coinmenced during early April. Puparia
Adults were particularly common in were formed within the rhizomes during
June. Eggs found on the surface of the last two weeks of April. Larvae and
Eleocharis stems had an incubation puparia were easily distinguished from
period of 5-7 days. Newly hatched lar- those of Chlorops and other species of
vae bored through the stem cuticle and Diplotoxa by their greenish color. The life
subsequently fed on the tissue within the cycle was elucidated by Wearsch and
stem. Larvae overwintered as first and Foote (1994) who also described and
second instars within stems, with the illustrated the egg and all larval instars.
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Table 1. Species, numbers collected, and trophic guilds ofhigher Diptera associated with Eleocharis
palustris in northeastern Ohio.
NumberofAdults
Species Collected TrophicGuild
Agromyzidae
Cerodontha dorsalts (Loew) 1 Leafminer ofgrasses
Undetermined sp. 2 Unknown
Anthomyzidae
Anthomyza sp. 1 Stem borer
Mumetopia occipitalis Melander 2 Stem borer
Chloropidae
Apallates neocoxendix (Sabrosky) 26 Unknown
Apallatesparticeps (Becker) 1 Unknown
Chlorops obscuricornis Loew 209 Stem borer ofEleocharis
Diplotoxa inclinata Becker 112 Stem borer ofEleocharis
Diplotoxa nigripes (Coquillett) 30 Stem borer ofEleocharis
Diplotoxa near versicolor 14 Stem borer ofEleocharis
Elliponeura debilis Loew 11 Seed predator ofEleocharis
Eribolus longulus (Loew) 1 Secondaryinvaderofdamagedstems
Incertella minor (Adams) 1 Secondaryinvaderofdamagedstems
Incertella incerta (Becker) 1 Secondaryinvaderofdamagedstems
Meromyza americana Fitch 1 Stem borer ofgrasses
Oscinellafrit Linneaus 4 Stem borer ofgrasses
Pseudopachychaeta approximatonervis 14 Seed predator ofEleocharis
(Zett.)
Rhopcdopterimi carbonaria (Loew) 51 Secondaryinvaderofdamaged stems
Thaumatomyiaglabra (Meigen) 1 Predator ofroot aphids
Drosophilidae
Scaptomyzapallida (Zetterstedt) 11 Scavenger ofdetritus
Ephydridae
Hydrelliaformosa Loew 2 Leafminer ofgrasses
Hydrellia griseola (Fallen) 1 Leafminer ofmany plant taxa
Leptopsilopa atrimana (Loew) 1 Scavenger ofdetritus
Notiphila caudata Fallen 44 Scavenger ofdetritus
Notiphila scalaris Loew 9 Scavenger ofanaerobic detritus
Ochthera anatolikos Clausen 2 Insect predator
Pelina truncatula Loew 1 Consumer ofCyanobacteria
Scatella stagnalis (Fallen) 10 Scavenger, phycovore
Typopsilopa atra (Loew) 12 Secondaryinvaderofdamagedstems
Milichiidae
Phyllomyza securicornis Fallen Unknown
Opomyzidae
Opomyzapetrel Mesnil Stem borer ofgrasses
Otitidae
Chaetopsis massyla (Walker) Secondaryinvaderofdamaged stems
Sciomyzidae
Atrichomelinapubera (Loew) 9 Predator ofstranded aquatic snails
Pherbellia nana (Fallen) 16 Predator ofstranded aquatic snails
Pherbelliaparallela (Walker) 4 Predator ofstranded aquatic snails
Pherbellia schoenherrimaculata (Cresson) 2 Predator ofamber snails
Dictya borealis Curran 1 Predator ofaquatic snails
Dictya expansa Steyskal 1 Predator ofaquatic snails
Dictya sabroskyi Steyskal 9 Predator ofaquatic snails
—
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Table 1. Continued.
Numberol'Adults
Spc Colkvleil Trophic(hiild
Diclya Mey.skci/i Valley 1 Predator ofaquatic snails
Sepedon armipc.s Loew 8 Predator ofaquatic snails
Sepedon fuscipcnnis Loew 28 Predator ofaquatic snails
Sepedon leniiieornis Cresson 5 Predator ofaquatic snails
Sepsidae
EnicoDiirci minor (Haliday) 5 Scavenger ofdetritus
Sepsispiinciuin (Fabricius) 6 Scavenger ofdetritus, feces
Sphaeroceridae
Leptocera sp. 39 Scavenger ofdetritus
Muscidae
Coenosia tigrina (Fabricius) 18 Insect predator
Lispa sp. 29 Insect predator
Schoenomyza chrysostoma Loew 21 Stem borer ofEleocharis
Schoenomvza dorsalis Loew 8 Stem borer ofEleocharis
Number ofspecies: 50.
Number ofgenera: 35.
Number ofindividuals: 875.
—
Diplotoxa nigripes (Coquillett). An early April. Puparia were formed within
uncommon species in the stand of E. the stems in early April, with emergence
paliistris, this small, dark species was occurring in early May. The wasp
common to abundant in nearby stands of Chaenusa sp. (Hymenoptera: Braconi-
the annual spike-rush, E. obtusa (Willd.) dae) was an important parasitoid, as 15
Scultes. The larvae are stem borers. The of 16 collected puparia were infested.
life cycle and the immature stages were The life cycle and immature stages were
described and illustrated by Wearsch and described and illustrated by Wearsch and
Foote (1994). Foote (1994).
—
Diplotoxa sp. near versicolor (Loew). Elliponeura debilis Loew. This was
The late Curtis W. Sabrosky (personal an uncommon, multivoltine seed preda-
communication) recognized this form as tor, having a flight period extending
a distinct species near D. versicolor, from mid-May to late September. Eggs
based upon the presence of white hairs were attached to the lower bracts sub-
on the dorsum ofthe abdomen instead of tending the inflorescence, with only 1 or
the dark hairs found in the D. versicolor. 2 eggs being found on each inflorescence.
It was a fairly common species in the Examination of inflorescence collected
stand ofEleocharis, whereas adults ofD. from the stand revealed infestation rates
versicolor itselfwere far more commonly varying from zero to 27%. Larvae fed on
encountered on stands of the path rush, the developing achenes, with each larva
Juncus tenuis Willd. (Juncaceae). This consuming 3-5 seeds. Puparia were
stem borer was an univoltine species in formed in the inflorescence within the
northeastern Ohio, with a flight period space formerly filled by an eaten achene.
extending from early May to late Sep- The pupal period lasted 6 8 days.
tember. Overwintering occurred as sec- Pseudopachychaeta approximatonervis
—
ond instars within the plant stems. Fully (Zetterstedt). Like E. debilis, this spe-
grown larvae were found near the bases cies fed on the developing achenes, and
of Eleocharis during late March and had a similar life cycle and flight period.
——
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109, 1 13
—
However, older larvae and puparia were Scatella stagnalis (Fallen). This was
yellowish in color, whereas those of E. a relatively common species in the
debilis, were greenish. Interestingly, only Eleocharis stand, with 10 specimens
one species ofthe two seed predators was being recorded. The larvae are general-
found in each infested inflorescence. An ized consumers of algae and organic
indication that this species is not strictly detritus (Zack and Foote 1978, Foote
a seed predator was the rearing ofadults 1979). —
from larvae that were feeding within Typopsilopa atra (Loew). This was
damaged stems of yellow nelumbo, a common species, becoming particularly
Nelumbo lutea (Willd.) Persoon (Nym- noticeable in mid-August. Its larvae are
phaeaceae), that had been attacked by saprophagous, having been reared in
larvae of the moth Bellura obliqua laboratory cultures of decaying lettuce
A
(Walker) (Noctuidae) in northern Ohio (Foote 1995). related species, T. nigra
(J. B. Keiper, personal communication). (Williston), was found to be a secondary
Rhopalopterum carbonaria (Loew). invader of the stems of California
This was a common species within the bulrush and cattail (Typha sp.) damaged
stand, with over 50 specimens being by the feeding of larvae of the primary
collected. Its larvae are secondary in- invader Bellura obliqua gargantua (Dyar)
vaders of culms of wetland monocots (Lepidoptera: Noctuidae) in southern
that previously had been damaged by California (Keiper et al. 2001).
primarily phytophagous species (Valley
Family Opomyzidae
et al. 1969). —
Opomyza petrel Mesnil. Although
Family Drosophilidae this was an abundant species in the
Eleocharis stand, occurring throughout
Scaptomyza pallida (Zetterstedt). the collecting season, it was not directly
This was a fairly common species within associated with the sedge. Its larvae are
the stand being found in low numbers known to be stem borers ofgrasses (Nye
throughout the summer months. Its 1958) which were abundant on either
larvae are saprophagous in decaying side of the Eleocharis stand. This is
plant material (Ferrar 1987). a common, non-native species in the
eastern states.
Family Ephydridae
— Family Otitidae
Notiphila candata Fallen. This was —
an abundant species during late May and Chaetopsis massyla (Walker). This
early June. Its larvae were non-selective was a relatively uncommon species with-
consumers of unicellular algae and de- in the Eleocharis stand. It appeared to be
caying particulate plant material (Eastin multivoltine, as adults were taken
and Foote 1971). throughout the summer. The larvae are
—
Notiphila scalaris Loew. A few speci- secondary invaders of the stems of
mens of this small species were taken wetland plants (Allen and Foote 1992),
within the Eleocharis stand, but it was far but no larvae were found in the stems of
more abundant in nearby stands of rice Eleocharis.
cutgrass. Keiper and Walton (2002)
collected numerous adults from a mono- Family Sciomyzidae
—
culture stand of California bulrush, Atrichomelina pubera (Loew). This
Schoenoplectus californicus (Meyer) So- species was regularly encountered in the
jak in California. Eleocharis stand, and appeared to be
——
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
14
multivoltine. Its larvae are generalized also prey on earthworms and other soft-
predators of aquatic pulmonale snails bodied invertebrates in moist to sodden
that have been stranded by dropping soils (Ferrar—1987).
water levels (Foote et al. 196—0). Lispe sp. Adults of this species were
Pherbellia nana (Fallen). This was found throughout the summer. The
a fairly common species, occurring in larvae are known to be general predators
low numbers throughout the summer of soft-bodied invertebrates (Skidmore
months. Like those of A. pubera, its 1985).
larvae prey on stranded aquatic snails Schoenomyza ch—rysostoma Loew, and
(Bratt et al. 1969). S. dorsalis Loew. Both Schoenomyza
Dictya expansa Steyskal, D. sabroskyi spp. were commonly taken in the stand,
Steyskal, and D. steyskali Valley. being particularly abundant in June.
Adults of these Dictya spp were taken Larvae are stem borers of wetland
in low numbers throughout the summer. monocots (Skidmore 1985, Ferrar 1987).
They are known to be multivoltine, and
Discussion
their larvae are generalized predators of
aquatic pulmonale snails (Valley and There was obvious partitioning of
Berg 1977). food resources within the stand of
Sepedon arinipes Loew, S. fuscipennis Eleocharis, and eight trophic guilds were
Loew, and S. tenuicornis Cresson. recognized (Table 1). A large guild con-
These Sepedon spp. were uncommon to sisting of 1 1 species fed as generalized
abundant within the stand ofEleocharis. scavengers of decaying organic matter.
Larvae of all three prey on aquatic The guild of secondary invaders con-
pulmonale snails (Neff and Berg 1966). sisted of six species whose larvae fed as
scavengers on stem tissues previously
Family Sepsidae damaged by phytophagous species. The
Enicomira minor (Haliday), Sepsis phytophagous community consisted of
—
piincta (Fabricius). Both of these spe- 1 1 species ofstem borers, three species of
cies were minor components of the fly leaf miners, two species of seed preda-
community, occurring in low numbers tors, and one species that fed on blue-
throughout the summer. Larvae of both green algae (Cyanobacteria). Predacious
are saprophagous on decaying plant species consisted of a snail-killing guild
material and feces (Ferrar 1987). that contained 1 1 species and a insect-
attacking guild that contained two spe-
Family Sphaeroceridae cies. The larval feeding habits of four
— species remain unknown. Only six ofthe
Leptocera spp.. Although numerous 50 species (12%) encountered within the
adults of this genus were swept from the stand fed directly on tissues of E.
stand, no larvae were found. Ferrar
pahistris, although another four species
(1987) reported that larvae are sapropha-
fed as secondary invaders of Eleocharis
gous, coprophagous, and necropha- stems that had been damaged by truly
gous.
phytophagous insects.
The family containing the greatest
Family Muscidae
— number of species within the stand was
Coenosia tigrina (Fabricius). This Chloropidae with 15. The same family
was a common species, occurring also produced the greatest number of
throughout the summer. Larvae are individuals (n = 477). Over 65% ofthese
predators of the stem-boring chloropid belonged to two species of stem borers,
larvae in sedges (Kuehne 1991). Larvae Chlorops obscuricornis (n = 209) and
VOLUME NUMBER
109, 1 15
Diplotoxa inclinata (n = 112). The . 1995. Biology of shore flies. Annual
abundance of these two species suggests Review ofEntomology 40: 417^42.
that larval infestations of the stems and . 2004a. AcalyptrateDipteraassociatedwith
stands of Carex lacustris and C. striata
rhizomes could be significant.
(Cyperaceae) in northeastern Ohio. Proceed-
The degree of possible competition ings ofthe Entomological Society ofWashing-
that exists among the six species that fed ton 106: 166-175.
as larvae within the Eleocharis stems is . 2004b. Acalyptrate Diptera ofa fen at the
unknown and warrants additional study. HerrickPreserveinPortageCounty, Ohio. The
Interactions among the two species of Great Lakes Entomologist 35: 131-148.
Foote, B. A., S. E. Neff, and C. O. Berg. 1960.
seed predators, E. debilis and P. approx- Biology and immature stages ofAtrichomelina
unatonervis, and species of other insect pubera (Diptera: Sciomyzidae). Annals of the
families (Curculionidae, Cecidomyiidae, Entomological Society of America 53:
undetermined family of Lepidoptera) 192-199.
that were also encountered in the in- Keiper, J. B. and W. E. Walton. 2002. Effects of
three management strategies on shore-flies
florescence need to be elucidated. (Diptera: Ephydridae) in newly constructed
treatment wetlands. Annals ofthe Entomolog-
Acknowledgments
ical Society ofAmerica 95: 570-576.
I am indebted to Barbara Andreas, Keiper, J. B., P. L. Brutsche, and B. A. Foote.
Kent State University, for aid in de- 1998. Acalyptrate Diptera associated with
water willow, Justicia americana (Acantha-
termining the species ofEleocharis at the ceae). Proceedings ofthe Entomological Soci-
study site. J. B. Keeper of the Cleveland ety ofWashington 100: 576-587.
Museum ofNatural History and Eric G. Keiper, J. B., J. Jiannino, M. Sanford, and W. E.
Chapman of Kent State University Walton. 2001. Biology and immature stages of
offered suggestions for the improvement Typopsilopa nigra (Williston) (Diptera: Ephy-
dridae), a secondary consumer of damaged
of the manuscript.
stemsofwetlandmonocots. Proceedings ofthe
Entomological Society of Washington 103:
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