New data indicate that larvae previously described as Spinadis Edmunds and Jensen and Spinadis wallacei Edmunds and Jensen are the larval stage of the prior named adults of Anepeorus simplex (Walsh) and are therefore synonymized. It is highly probable that larvae described as Acanthomola pubescens Whiting and Lehmkuhl represent the larval stage of Anepeorus rusticus McDunnough, but they remain to be reared. Anepeorus sensu novum and Pseudiron are placed in the subfamily Anepeorinae, and Raptoheptagenia (= recently associated Heptagenia cruentata Walsh adults and larvae previously thought to be Anepeorus) is placed in the subfamily Heptageniinae

McCafferty, W. P

Provonsha, A. V

English

ValpoScholar

The Great Lakes Entomologist 
Volume 21 
Number 1 - Spring 1988 Number 1 - Spring 1988 Article 4 
April 1988 
Revisionary Notes on Predaceous Heptageniidae Based on Larval 
and Adult Associations (Ephemeroptera) 
W. P. McCafferty 
Purdue University 
A. V. Provonsha 
Purdue University 
Follow this and additional works at: https://scholar.valpo.edu/tgle 
 Part of the Entomology Commons 
Recommended Citation 
McCafferty, W. P. and Provonsha, A. V. 1988. "Revisionary Notes on Predaceous Heptageniidae Based on 
Larval and Adult Associations (Ephemeroptera)," The Great Lakes Entomologist, vol 21 (1) 
Available at: https://scholar.valpo.edu/tgle/vol21/iss1/4 
This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. 
It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. 
For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 
1988 THE GREAT LAKES ENTOMOLOGIST 15 
REVISIONARY NOTES ON PREDACEOUS HEPTAGENIIDAE BASED 
ON 
LARVAL AND ADULT ASSOCIATIONS (EPHEMEROPTERA) 
W. P. McCafferty and A. V. Provonsha1 
ABSTRACT 
New data indicate that larvae previously described as Spinadis Edmunds and Jensen and 
Spinadis wallacei Edmunds and Jensen are the larval stage of the prior named adults of 
Anepeorus simplex (Walsh) and are therefore synonymized. It is highly probable that 
larvae described as Acanthomola pubescens Whiting and Lehmkuhl represent the larval 
stage of Anepeorus rusticus McDunnough, but they remain to be reared. Anepeorus sensu 
novum and Pseudiron are placed in the subfamily Anepeorinae, and Raptoheptagenia (= 
recently associated Heptagenia cruentata Walsh adults and larvae previously thought to 
be Anepeorus) is placed in the subfamily Heptageniinae. 
Recent studies by McCafferty and Provonsha (1984, 1985, 1986) dealt with the 
predaceous heptageniid mayflies that have been traditionally placed in the genera 
Anepeorus McDunnough, Pseudiron McDunnough, and Spinadis Edmunds and Jensen. 
Questions were raised with regard to certain larval associations and application of generic 
nomenclature to the larval stage. The nomenclatural propriety of Pseudiron, which was 
recently revised by Pescador (1985), is not in doubt. The larva first assigned to Anepeorus 
by Burks (1953) was clearly stated to be questionable and provisional, but by the 
appearance of the treatment of North American mayflies by Edmunds et al. (1976), this 
form had been generally accepted without question as correct, although such larvae had 
never been reared, and McCafferty and Provonsha (1985, 1986) were highly suspect of 
the association. The genus Spinadis was based on a very distinctive larval form (Edmunds 
and Jensen 1974), but the fact that only a female adult has been reared from such larvae 
has not clearly resolved the question of whether its adult stage had been previously 
described under a different name (McCafferty and Provonsha 1984). 
Two recent findings lead us to now suppress the name Spinadis. Reexamination of the 
female adult (McCafferty and Provonsha 1984) reared from a larva referable to Spinadis 
indicates that it is similar to adult females tentatively assigned to Anepeorus that were 
taken in series with male adults of Anepeorus simplex (Walsh). Similarities include wing 
venation, shape of the costal projection of the hindwing, hind tarsus to tibia ratio, ventral 
margin of the frontal shelf of the head, margination of the eyes dorsally, and elongation 
and posterior margin of the pronotum. These morphological traits (see McCafferty and 
Provonsha [1984] for details) are not only strikingly similar in these females but also very 
similar in males that are without doubt Anepeorus. 
In addition, larvae referable to Anepeorus were very recently reared in Saskatchewan 
to adults of Heptagenia cruentata Walsh by Whiting and Lehmkuhl (1987a), who erected 
a new genus, Raptoheptagenia, for that species. Thus, with the accurate placement of 
those larvae, there remains no contradiction that larvae originally named Spinadis are the 
IDepartment of Entomology, Purdue University, Wesl Lafayette, IN 47907. 
1
McCafferty and Provonsha: Revisionary Notes on Predaceous Heptageniidae Based on Larval and
Published by ValpoScholar, 1988
16 THE GREAT LAKES ENTOMOLOGIST Vol. 21, No. 1 
larvae of Anepeorus simplex. Also, known distributional and large-river habitat data for 
larvae known as Spinadis and adults of A. simplex in eastern North America are similar. 
In consideration of the above we designate Spinadis Edmunds and Jensen as a NEW 
JUNIOR SYNONYM of Anepeorus McDunnough, and Spinadis wallacei Edmunds and 
Jensen as a NEW JUNIOR SYNONYM of Anepeorus simplex (Walsh). The genus 
Anepeorus is very distinctive in both the adult stage (McCafferty and Provonsha 1985) 
and larval stage (described as Spinadis [Edmunds et at. 1976, McCafferty and Provonsha 
1986]). 
The larval stage of Anepeorus rusticus McDunnough, the western North American 
species, remains unknown. Adults of this species are clearly congeneric with, but 
distinguishable from, those of A. simplex (McCafferty and Provonsha 1985), but it is 
problematic as to how similar the larvae of the two species will prove to be. We predict 
that A. rusticus larvae will be a two-tailed predaceous form (with reduced mouthpart 
setation and reduced molar region) with gill structure similar to A. simplex. The presence 
of 
spines, or tubercles, dorsally on the body 
ofA. simplex may be of generic significance. 
The head shape of A. rusticus larvae could be somewhat different than that of A. simplex 
since there are distinctive differences in the frontal shelf of the adults of the two species 
(McCafferty and Provonsha 1985). 
In consideration f all of the bove, it appears extremely probable that the larvae 
recently described as Acamhomola pubescens Whiting and Lehmkuhl (1987b) from 
Saskatchewan is actually the larval stage of Anepeorus rusticus. The similarity in 
characteristics to those of larval A. simplex is comparable in degree to the similarities 
between the adults of he two Anepeorus species. The range and general habitat for A. 
rusticus adults include that of the newly described larvae. Although it is our opinion that 
Acanthomola pubescens is synonymous with Anepeorus rusticus, it would appear prudent 
to defer formal synonymy until verified by rearing, especially in light of the recent history 
of 
misassociations within the predaceous Heptageniidae. Unfortunately, Whiting and 
Lehmkuhl (1987b) stated that because 
of its rarity and threats to its habitat the larvae 
probably would never be reared. 
Spinadis and Pseudiron were found to be closely related based on larval mouthparts 
(McCafferty and Provonsha 1986). We now consider Anepeorus sensu novum and 
Pseudiron to be sister genera for the same reasons, and because of additional adult 
synapomorphies. We therefore place Ps udiron and Anepeorus (including nominal 
Acanthomola) together in the subfamily Anepeorinae rather than in separate subfamilies 
as 
has been done 
in the past. 
Raptoheptagenia is more plesiomorphic than Anepeorus and Pseudiron, and it appears 
to 
be either independently derived or intermediate between the more generalized 
heptageniid genera traditionally placed 
in the subfamily Heptageniinae and the specialized 
Anepeorus and Pseudiron. The larvae of Raptoheptagenia, although predaceous and with 
a highly modified gill structure, have a body form more typical of Heptageniinae. They 
also possess a number of autapomorphies, such a  raptorial labial palps (McCafferty and 
Provonsha 1986). The adult of Raptoheptagenia has remained a more generalized form 
typical of many Heptageniinae. McDunnough (1931), however, noted certain similarities 
between the adults of Pseudiron and Raptoheptagenia (as Heptagenia cruentata). We 
prefer to include Raptoheptagenia in the subfamily Heptageniinae until its relationships 
can be more fully elucidated. 
ACKNOWLEDGMENTS 
This paper is published as Purdue Experiment Station Journal No. 11 ,205. 
2
The Great Lakes Entomologist, Vol. 21, No. 1 [1988], Art. 4
https://scholar.valpo.edu/tgle/vol21/iss1/4
1988 THE GREAT LAKES ENTOMOLOGIST 17 
LITERATURE CITED 
Burks, B. D. 1953. The mayflies, or Ephemeroptera, of Illinois. BulL Illinois Nat. His!. Surv. 
26:1-216. 
Edmunds, G. F., Jr., and S. L. Jensen. 1974. A new genus and subfamily of North American 
Heptageniidae (Ephemeroptera). Proc. EntomoL Soc. Washington 76:495-497. 
Edmunds, G. F., Jr., S. L. Jensen, and L. Berner. 1976. The mayflies of North and Central 
America. Univ. Minnesota Press, Minneapolis, 330 pp. 
McCafferty, W. P., and A. V. Provonsha. 1984. The first adult of Spinadis (Ephemeroptera: 
Heptageniidae). EntomoL News 95:173-179. 
1985. Systematics of Anepeorus (Ephemeroptera: Heptageniidae). Great Lakes EntomoL 
18:1-6. 
1986. Comparative mouthpart morphology and evolution ofthe carnivorous Heptageniidae 
(Ephemeroptera). Aq. Ins. 8:83-89. 
McDunnough, J. 1931. New species of North American Ephemeroptera. Canadian EntomoL 
63:82-93. 
Pescador, M. L. 1985. Systematics of the Nearctic genus Pseudiron (Ephemeroptera: Heptageniidae: 
Pseudironinae). Florida Entomol. 68:432-444. 
Whiting, E. R., and D. M. lehmkuhL 1987a. Raptoheptagenia cruentata, gen. nov. (Ephemerop­
lera: Heptageniidae), new association of the larva previously thought to be AnepeorU8 with the 
adult of Heptagenia cruentata Walsh. Canadian EntomoL 119:405-407. 
___
. 1987b. 
Acanthomola pubescens, a new genus and species of Heptageniidae (Ephemcrop­
tera) from western Canada. Canadian EntomoL 119:409-417. 
3
McCafferty and Provonsha: Revisionary Notes on Predaceous Heptageniidae Based on Larval and
Published by ValpoScholar, 1988


Revisionary Notes on Predaceous Heptageniidae Based on Larval and Adult Associations (Ephemeroptera)

W. P McCafferty

A. V Provonsha

Crossref

Valparaiso University

The Great Lakes Entomologist Volume 21 Number 1 - Spring 1988 Number 1 - Spring 1988 Article 4 April 1988 Revisionary Notes on Predaceous Heptageniidae Based on Larval and Adult Associations (Ephemeroptera) W. P. McCafferty Purdue University A. V. Provonsha Purdue University Follow this and additional works at: https://scholar.valpo.edu/tgle  Part of the Entomology Commons Recommended Citation McCafferty, W. P. and Provonsha, A. V. 1988. "Revisionary Notes on Predaceous Heptageniidae Based on Larval and Adult Associations (Ephemeroptera)," The Great Lakes Entomologist, vol 21 (1) Available at: https://scholar.valpo.edu/tgle/vol21/iss1/4 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 1988 THE GREAT LAKES ENTOMOLOGIST 15 REVISIONARY NOTES ON PREDACEOUS HEPTAGENIIDAE BASED ON LARVAL AND ADULT ASSOCIATIONS (EPHEMEROPTERA) W. P. McCafferty and A. V. Provonsha1 ABSTRACT New data indicate that larvae previously described as Spinadis Edmunds and Jensen and Spinadis wallacei Edmunds and Jensen are the larval stage of the prior named adults of Anepeorus simplex (Walsh) and are therefore synonymized. It is highly probable that larvae described as Acanthomola pubescens Whiting and Lehmkuhl represent the larval stage of Anepeorus rusticus McDunnough, but they remain to be reared. Anepeorus sensu novum and Pseudiron are placed in the subfamily Anepeorinae, and Raptoheptagenia (= recently associated Heptagenia cruentata Walsh adults and larvae previously thought to be Anepeorus) is placed in the subfamily Heptageniinae. Recent studies by McCafferty and Provonsha (1984, 1985, 1986) dealt with the predaceous heptageniid mayflies that have been traditionally placed in the genera Anepeorus McDunnough, Pseudiron McDunnough, and Spinadis Edmunds and Jensen. Questions were raised with regard to certain larval associations and application of generic nomenclature to the larval stage. The nomenclatural propriety of Pseudiron, which was recently revised by Pescador (1985), is not in doubt. The larva first assigned to Anepeorus by Burks (1953) was clearly stated to be questionable and provisional, but by the appearance of the treatment of North American mayflies by Edmunds et al. (1976), this form had been generally accepted without question as correct, although such larvae had never been reared, and McCafferty and Provonsha (1985, 1986) were highly suspect of the association. The genus Spinadis was based on a very distinctive larval form (Edmunds and Jensen 1974), but the fact that only a female adult has been reared from such larvae has not clearly resolved the question of whether its adult stage had been previously described under a different name (McCafferty and Provonsha 1984). Two recent findings lead us to now suppress the name Spinadis. Reexamination of the female adult (McCafferty and Provonsha 1984) reared from a larva referable to Spinadis indicates that it is similar to adult females tentatively assigned to Anepeorus that were taken in series with male adults of Anepeorus simplex (Walsh). Similarities include wing venation, shape of the costal projection of the hindwing, hind tarsus to tibia ratio, ventral margin of the frontal shelf of the head, margination of the eyes dorsally, and elongation and posterior margin of the pronotum. These morphological traits (see McCafferty and Provonsha [1984] for details) are not only strikingly similar in these females but also very similar in males that are without doubt Anepeorus. In addition, larvae referable to Anepeorus were very recently reared in Saskatchewan to adults of Heptagenia cruentata Walsh by Whiting and Lehmkuhl (1987a), who erected a new genus, Raptoheptagenia, for that species. Thus, with the accurate placement of those larvae, there remains no contradiction that larvae originally named Spinadis are the IDepartment of Entomology, Purdue University, Wesl Lafayette, IN 47907. 1McCafferty and Provonsha: Revisionary Notes on Predaceous Heptageniidae Based on Larval andPublished by ValpoScholar, 198816 THE GREAT LAKES ENTOMOLOGIST Vol. 21, No. 1 larvae of Anepeorus simplex. Also, known distributional and large-river habitat data for larvae known as Spinadis and adults of A. simplex in eastern North America are similar. In consideration of the above we designate Spinadis Edmunds and Jensen as a NEW JUNIOR SYNONYM of Anepeorus McDunnough, and Spinadis wallacei Edmunds and Jensen as a NEW JUNIOR SYNONYM of Anepeorus simplex (Walsh). The genus Anepeorus is very distinctive in both the adult stage (McCafferty and Provonsha 1985) and larval stage (described as Spinadis [Edmunds et at. 1976, McCafferty and Provonsha 1986]). The larval stage of Anepeorus rusticus McDunnough, the western North American species, remains unknown. Adults of this species are clearly congeneric with, but distinguishable from, those of A. simplex (McCafferty and Provonsha 1985), but it is problematic as to how similar the larvae of the two species will prove to be. We predict that A. rusticus larvae will be a two-tailed predaceous form (with reduced mouthpart setation and reduced molar region) with gill structure similar to A. simplex. The presence of spines, or tubercles, dorsally on the body ofA. simplex may be of generic significance. The head shape of A. rusticus larvae could be somewhat different than that of A. simplex since there are distinctive differences in the frontal shelf of the adults of the two species (McCafferty and Provonsha 1985). In consideration f all of the bove, it appears extremely probable that the larvae recently described as Acamhomola pubescens Whiting and Lehmkuhl (1987b) from Saskatchewan is actually the larval stage of Anepeorus rusticus. The similarity in characteristics to those of larval A. simplex is comparable in degree to the similarities between the adults of he two Anepeorus species. The range and general habitat for A. rusticus adults include that of the newly described larvae. Although it is our opinion that Acanthomola pubescens is synonymous with Anepeorus rusticus, it would appear prudent to defer formal synonymy until verified by rearing, especially in light of the recent history of misassociations within the predaceous Heptageniidae. Unfortunately, Whiting and Lehmkuhl (1987b) stated that because of its rarity and threats to its habitat the larvae probably would never be reared. Spinadis and Pseudiron were found to be closely related based on larval mouthparts (McCafferty and Provonsha 1986). We now consider Anepeorus sensu novum and Pseudiron to be sister genera for the same reasons, and because of additional adult synapomorphies. We therefore place Ps udiron and Anepeorus (including nominal Acanthomola) together in the subfamily Anepeorinae rather than in separate subfamilies as has been done in the past. Raptoheptagenia is more plesiomorphic than Anepeorus and Pseudiron, and it appears to be either independently derived or intermediate between the more generalized heptageniid genera traditionally placed in the subfamily Heptageniinae and the specialized Anepeorus and Pseudiron. The larvae of Raptoheptagenia, although predaceous and with a highly modified gill structure, have a body form more typical of Heptageniinae. They also possess a number of autapomorphies, such a  raptorial labial palps (McCafferty and Provonsha 1986). The adult of Raptoheptagenia has remained a more generalized form typical of many Heptageniinae. McDunnough (1931), however, noted certain similarities between the adults of Pseudiron and Raptoheptagenia (as Heptagenia cruentata). We prefer to include Raptoheptagenia in the subfamily Heptageniinae until its relationships can be more fully elucidated. ACKNOWLEDGMENTS This paper is published as Purdue Experiment Station Journal No. 11 ,205. 2The Great Lakes Entomologist, Vol. 21, No. 1 [1988], Art. 4https://scholar.valpo.edu/tgle/vol21/iss1/41988 THE GREAT LAKES ENTOMOLOGIST 17 LITERATURE CITED Burks, B. D. 1953. The mayflies, or Ephemeroptera, of Illinois. BulL Illinois Nat. His!. Surv. 26:1-216. Edmunds, G. F., Jr., and S. L. Jensen. 1974. A new genus and subfamily of North American Heptageniidae (Ephemeroptera). Proc. EntomoL Soc. Washington 76:495-497. Edmunds, G. F., Jr., S. L. Jensen, and L. Berner. 1976. The mayflies of North and Central America. Univ. Minnesota Press, Minneapolis, 330 pp. McCafferty, W. P., and A. V. Provonsha. 1984. The first adult of Spinadis (Ephemeroptera: Heptageniidae). EntomoL News 95:173-179. 1985. Systematics of Anepeorus (Ephemeroptera: Heptageniidae). Great Lakes EntomoL 18:1-6. 1986. Comparative mouthpart morphology and evolution ofthe carnivorous Heptageniidae (Ephemeroptera). Aq. Ins. 8:83-89. McDunnough, J. 1931. New species of North American Ephemeroptera. Canadian EntomoL 63:82-93. Pescador, M. L. 1985. Systematics of the Nearctic genus Pseudiron (Ephemeroptera: Heptageniidae: Pseudironinae). Florida Entomol. 68:432-444. Whiting, E. R., and D. M. lehmkuhL 1987a. Raptoheptagenia cruentata, gen. nov. (Ephemerop­lera: Heptageniidae), new association of the larva previously thought to be AnepeorU8 with the adult of Heptagenia cruentata Walsh. Canadian EntomoL 119:405-407. ___. 1987b. Acanthomola pubescens, a new genus and species of Heptageniidae (Ephemcrop­tera) from western Canada. Canadian EntomoL 119:409-417. 3McCafferty and Provonsha: Revisionary Notes on Predaceous Heptageniidae Based on Larval andPublished by ValpoScholar, 1988

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Revisionary Notes on Predaceous Heptageniidae Based on Larval and Adult Associations (Ephemeroptera)

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