Nelson, C.R., R.W. Baumann, & J.J. Lee. 2013. New morphological observations and phylogenetic placement of Capnia
shasta (Plecoptera: Capniidae). Illiesia, 9(12):122-125. Available online: http://www2.pms-lj.si/illiesia/nliesia09-12.pdf
http://zoobank.Org/urn:lsid:zoobank.org:pub:5F49A455-D4A6-4DAD-9E29-393188457860
NEW MORPHOLOGICAL OBSERVATIONS AND PHYLOGENETIC PLACEMENT OF CAPNIA SHASTA (PLECOPTERA: CAPNIIDAE)
C. Riley Nelson1, Richard W. Baumann1 and Jonathan J. Lee2
1 Department of Biology and Monte L. Bean Life Science Museum, Brigham Young University,
Provo, Utah, 84602 U.S.A. E-mail: rileynelson@byu.edu; E-mail: richard_baumann@byu.edu
2 2337 15th Street, Eureka, California, 95501 U.S.A. E-mail: jlee@humboldt1.com
ABSTRACT
The male epiproct of Capnia shasta is redescribed and the presence of a lower lobe is noted. Capnia shasta is moved to the C. ventura species Subgroup of the C. californica Group as defined by Nelson (2004). A redescription of the male epiproct is provided and supported by scanning electron micrographs. We also report this species from the Salmon River drainage, Marble Mountains, northern California.
Keywords: Plecoptera, stonefly, Capniidae, Capnia, California, Coast Range
INTRODUCTION
Capnia shasta Nelson and Baumann (2009) has had an interesting history, appearing in the stonefly literature tangentially 24 years ago. It first appeared as a group of four illustrations incorrectly labeled as C. umpqua in Nelson and Baumann (1989) as was noted in Baumann and Stewart (2009) and noted and corrected in Nelson and Baumann (2009). Nelson and Baumann (2009) attempted to clarify the status of this species illustrated and keyed in Nelson and Baumann (1989) by describing C. shasta. Their description stated that the epiproct was composed of only a single well-developed dorsal process. This decision was based on the illustrations in Nelson and Baumann (1989) as well as the specimens and the scanning electron micrographs (SEM) in Nelson and Baumann (2009). Recently, one of the authors (J.J. Lee) collected specimens from near Sulphur Creek,
close to the type locality and additional specimens from a site along Salmon River Road in Siskiyou County, California. Male specimens clearly possessed a lower lobe on the epiproct. It was difficult to directly observe this structure because it was closely appressed to the abdomen beneath the larger upper lobe of the epiproct. Upon reexamining the male holotype of C. shasta we concluded that the lower process was missed in the original description, where the epiproct was characterized as "with only a single process well-developed" (Nelson and Baumann 2009). The epiproct of this species does indeed possess a lower process. The process is relatively small, usually closely appressed to, and sometimes appearing to be imbedded in, the softer dorsal surface of tergum 10. We provide the following addendum to the description of the epiproct.
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Nelson, C.R., R.W. Baumann, & J.J. Lee. 2013. New morphological observations and phylogenetic placement of Capnia shasta (Plecoptera:
Capniidae). Illiesia, 9(12):122-125. Available online: http://www2.pms-lj.si/illiesia/nliesia09-12.pdf
Figs. 1-8. Capnia shasta male genitalia. Figs. 1-4. California, Shasta County, creek near Castle Crags State Park. 1. Terminalia, dorsal. 2. Epiproct, dorsal. 3. Epiproct, dorsolateral. 4. Epiproct, lateral. Figs. 5-8. California, Siskiyou County, tributary, Salmon River, Salmon River Road. 5. Terminalia, dorsal. 6. Epiproct, dorsal. 7. Epiproct, dorsolateral. 8. Epiproct, dorsolateral, anterior.
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Volume 9 - Number 12 - Page 123
Nelson, C.R., R.W. Baumann, & J.J. Lee. 2013. New morphological observations and phylogenetic placement of Capnia shasta (Plecoptera:
Capniidae). Illiesia, 9(12):122-125. Available online: http://www2.pms-lj.si/illiesia/Illiesia09-12.pdf
MATERIALS AND METHODS
We examined specimens with Wild M-8 and Olympus SZX12 stereomicroscopes. Additional SEMs were made at the BYU Electron Optics Laboratory using a Philips XL30, ESEM FEG microscope as in Nelson and Baumann (2009). Figure plates were prepared at the M.L. Bean Museum graphics laboratory. Specimens studied are deposited at the following collections; Brigham Young University Collection, Provo, Utah (BYUC); Jon J. Lee Collection, Eureka, California (JJLC); United States National Collection, Smithsonian Institution, Washington, D.C. (USNM).
RESULTS AND DISCUSSION
Capnia shasta Nelson and Baumann
(Figs. 1-8)
Capnia umpqua Nelson and Baumann 1989:306. Capnia shasta Nelson and Baumann 2009:188. Type locality, California: Shasta County, Sulphur Creek, Castle Crags State Park, N 41.15587° W 122.36285°.
Material examined. California: Shasta Co., small tributary of Castle Creek flowing from north, above Castle Creek Road, approximately milepost 1.5, N 41.15614° W 122.35319°, 18 February 2010, J.J. Lee, 4$, 7? (BYUC, JJLC); Sulphur Creek, Castle Crags State Park, N 41.15587° W 122.36285°, 16 February 1985, R.W. Baumann & C.R. Nelson, holotype $ (USNM) and 3$, 4? paratypes (BYUC). Siskiyou Co., creek, mile 1.9, Salmon River Road, N 41.38061° W 123.46496°, 4 February 2011, J.J. Lee, 3$, 4? (BYUC, JJLC); 9 February 2011, J.J. Lee, 26$, 12? (BYUC, JJLC).
Male. Epiproct with inflated dorsal process and shorter ventral process. Other features as in original description by Nelson & Baumann (2009). When fully visible anteriorly, the epiproct has the appearance of a small bird with its gape wide open (Figs. 1-8). Discussion. Nelson (2004) included four species of Capnia in the C. ventura species Subgroup of the C. californica Group: Capnia kersti Nelson, C. regilla Nelson and Baumann, C. saratoga Nelson and Baumann, and C. ventura Nelson and Baumann. We are including C. shasta in the C. ventura Subgroup based on the epiproct structure. Capnia shasta can be
distinguished from the other subgroup members by features given in Nelson and Baumann (2009) and by the short lower process of the epiproct described herein.
Phylogeny. The cladogram given as Fig. 10 in Nelson and Baumann (2009) showed C. shasta as part of a polytomy with members of the C. californica Group. This hypothesis needs to be re-evaluated based on the actual state of the epiproct of C. shasta presented herein. With the recognition of the lower process of the epiproct, three characters in the matrix of Nelson and Baumann (2009) must be recoded: character 5 changes from 1 to 0 by redefining the appropriate upper process; 17 changes from 0 to 1 indicating an apomorphic deep notch between the two processes of the epiproct; and 18 changes to an apomorphic narrow lower process. In using the branch rearrangement facility in MacClade 4.08a (Maddison and Maddison 2005), two most parsimonious trees result, both of which clearly move C. shasta into the C. ventura Subgroup and require little change to the Nelson and Baumann (2009) hypothesis of relationships. One places C. shasta basal to all members of the subgroup (from C. regilla to C. ventura in Fig. 10 of Nelson and Baumann (2009)) and the other places this taxon further up the tree between C. regilla and C. saratoga. Analysis using new information we present here resolved the tree to a more basal polytomy and by consensus created a polytomy further up the tree, within the C. ventura Subgroup. Resolution of this polytomy will require using additional morphological or molecular characters.
Distribution. The most recent collections of C. shasta were made in small creeks that the authors predict have reduced surface flow in the summer. This species is now known from three localities in the Coast Ranges of northern California, two near Castle Crags and one in the Salmon River drainage.
ACKNOWLEDGMENTS
The holotype male of C. shasta was loaned by Oliver S. Flint, Jr., United States National Museum, Smithsonian Institution, Washington D.C. Scanning electron micrographs were produced with the help of Michael Standing, Electron Optics Laboratory, Brigham Young University. The figure plate was produced by Randal Baker, Graphic Artist, Monte L.
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Volume 9 - Number 12 - Page 124
Nelson, C.R., R.W. Baumann, & J.J. Lee. 2013. New morphological observations and phylogenetic placement of Capnia shasta (Plecoptera:
Capniidae). Illiesia, 9(12):122-125. Available online: http://www2.pms-lj.si/illiesia/Illiesia09-12.pdf
Bean Life Science Museum, Brigham Young
University.
REFERENCES
Baumann, R.W. and K.W. Stewart. 2009. What is Capnia umpqua Frison? (Plecoptera: Capniidae), distribution and variation of terminalia. Illiesia, 5 (5):34-39.
Maddison, D.R. and W.P. Maddison, 2005. MacClade 4: Analysis of phylogeny and character evolution. Version 4.08a. http://macclade.org.
Nelson, C.R. 2004. Systematics of the Capnia californica species group, including a morphological phylogeny, zoogeography, and description of Capnia kersti, new species (Plecoptera: Capniidae). Annals of the Entomological Society of America, 97 (1):97-104.
Nelson, C.R. and R.W. Baumann. 1989. Systematics and distribution of the winter stonefly genus Capnia (Plecoptera: Capniidae) in North America. Great Basin Naturalist, 49: 289-363 +3 page index.
Nelson, C.R. and R.W. Baumann. 2009. Capnia shasta, a new species in the californica group from northwestern California (Plecoptera: Capniidae). Illiesia, 5 (18):188-194.
Received 12 September 2013, Accepted 3 October 2013, Published 13 November 2013
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