A Review of the North American Freshwater Snail Genus Pyrgulopsis

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82 FIGURE 31.—Strict consensus cladogram for Pyrgulopsis species (ci = 0.24, ri = 0.57). SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY outgroup carinifera nanus fairbanksensis isolata bryantwalkeri conica manantiali micrococcus morrisoni simplex stearnsiana thompsoni arga lustrica ogmorphaphe pachyta letsoni scalariformis agarhecta castor halcyon arnargosae bacchus sola brandi bruneauensis owensensis perturbata aardahli giulianii montezumensis glandulosa robusta pecosensis hendersoni idahoensis bernardina avernalis crystalis arizonae erythropoma pisteri archimedis chihuahua intermedia - kolobensis pilsbryana trivialis roswellensis merriami gilae deserta chupaderae neomexicana davisi metcalfi californiensis wongi longinqua thermalis
NUMBER 554 83 and southeastern Arizona form a clade (Clade 3) defined by three transformations, including unique synapomorphies involving loss of penial lobe (30-3) and terminal gland (32-3). Other unique synapomorphies defining sub-clades within this group are an enlarged ventral gland (43-2) and superficial position of ventral gland (44-1). Members of this clade usually have globose shells, and their penes are ornamented by a ventral gland, sometimes accompanied by a large dorsal gland. A second western clade (Clade 4) is comprised of 10 species from eastern California, northern Arizona, and Snake River environs of Wyoming, Idaho, and Oregon; and includes all members of the subgenus Natricola sensu Gregg and Taylor (1965). This group is defined by five synapomorphies, of which only one, presence of a dot-like Dgl penial gland, is unique; this character is transformed in advanced members of this clade. Other defining characters are homplasious, and include a strong ventral callus on the operculum (10-1), and three features relating to Dgl (37-1, 38-2, 40-1). There is extensive parallelism in penial and female genitalic character transformations between this clade and that discussed below. Members of this group often are large, have ovate-conic shells, and have penes ornamented by terminal gland, Dgl, and sometimes Dg2 and/or Dg3. A third western clade (Clade 5), comprised of 18 species from various drainages, closely approximating the "Ffontelicella]. californiensis series" of Taylor (1987), is defined by five character-state transformations. Three of these (33-1,34-1, 35-1) are associated with the unique occurrence of a penial gland (Pg) in this clade. The other two, involving a strong ventral callus on operculum (10-1) and loss of anterior capsule gland vestibule (48-0), are homoplasious. Within this group, sub-clades are defined by die following unique or near-unique character-state transformations: proximally bifurcate penial gland (35-2, one parallel within this clade), Dgl a short strip (36-2), multiple minor dorsal glands (42-2, paralleled by two species), posterior position of ovary (46-1, parallel by one species within this clade), multiple ventral glands (43-2), and elongate bursa copulatrix (52-1, two parallels and one reversal). Members of this group have globose to elongate-conic shells and penes ornamented by a terminal and penial gland, often accompanied by one or more dorsal glands. The 17 Western American species having a relatively simple penis (ornamented solely by terminal gland) do not comprise a monophyletic group and, in general, are poorly resolved on the cladogram. Four globose-shelled species from southern Nevada (carinifera, nanus, fairbanksensis, isolata) are basally placed on the cladogram, but are not a well-defined group; and an assemblage of eight species scattered throughout the West, approximating the "F[ontelicellaJ. stearnsiana series" of Taylor, 1987), forms an unsupported polytomy. On the other hand, two small groups of species are more strongly supported. One pair from Gila River drainage of Arizona (bacchus, sola) is defined by a small penis (26-1, unique synapomorphy), bifurcate penial lobe (paralleled in one species), and simple terminal gland on penis (32-0, a homoplasious reversal). A second group, comprised of three highly disjunct species from internal drainage of northern Mexico (brandi), Amargosa River drainage (amargosae) and lower Snake River drainage (bruneauensis); is defined by three character-state transformations: weak ventral operculum attachment scar (9-0, homoplasious), penial filament (28-1, paralleled in one species), and bursal duct positioned lateral to albumen gland (59-0, homoplasious).
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A Review of the North American Fres
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S M I T H S O N I A N C O N T R I B
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Contents Page Introduction 1 Materi
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A Review of the North American Fres
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NUMBER 554 position of coiled ovidu
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NUMBER 554 field work and laborator
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NUMBER 554 FIGURE 2.—Morphology o
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NUMBER 554 Sr Ag Dsr FIGURE 4.—Di
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NUMBER 554 11 studied the anatomy o
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NUMBER 554 13 contrasted the genus
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NUMBER 554 15 FIGURE 7.—Western P
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NUMBER 554 17 Pyrgulopsis amargosae
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NUMBER 554 19 (in T 3S, R 22E, 25,0
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NUMBER 554 21 Pratt (1977) recorded
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NUMBER 554 23 DISTRIBUTION.—This
Page 31 and 32:
NUMBER 554 25 nucleus slightly ecce
Page 33 and 34:
NUMBER 554 27 FIGURE 11.—Western
Page 35 and 36:
NUMBER 554 29 FIGURE 12.—Western
Page 37 and 38: NUMBER 554 31 FIGURE 13.—Western
Page 39 and 40: NUMBER 554 33 ornament an elongate,
Page 41 and 42: NUMBER 554 35 dium centered slightl
Page 43 and 44: NUMBER 554 FIGURE 15.—Western Pyr
Page 45 and 46: NUMBER 554 Pyrgulopsis glandulosa H
Page 47 and 48: NUMBER 554 41 visceral coil black.
Page 53 and 54: NUMBER 554 copulatrix. Seminal rece
Page 55 and 56: NUMBER 554 Pyrgulopsis merriami (Pi
Page 59 and 60: NUMBER 554 or lightly pigmented on
Page 61 and 62: NUMBER 554 weakly frilled. Attachme
Page 63 and 64: NUMBER 554 lake have not yielded li
Page 65 and 66: NUMBER 554 59 REMARKS.—The small
Page 69 and 70: NUMBER 554 63 units; Dg3 small (som
Page 71 and 72: NUMBER 554 65 Pyrgulopsis simplex H
Page 75 and 76: NUMBER 554 69 DIAGNOSIS.—Shell ov
Page 77 and 78: NUMBER 554 71 filament, extending s
Page 79 and 80: NUMBER 554 FIGURE 28.—Eastern Pyr
Page 81 and 82: NUMBER 554 75 medium length, obliqu
Page 83 and 84: NUMBER 554 77 2e). Visceral coil li
Page 85 and 86: NUMBER 554 FIGURE 30.—Eastern Pyr
Page 87: NUMBER 554 ANSP 61606; paralectotyp
Page 91 and 92: Appendix 2 Fossil Species Described
Page 93 and 94: NUMBER 554 87 Pyrgulopsis bryantwal
Page 95 and 96: NUMBER 554 89 FIGURE 32.—Central
Page 97 and 98: NUMBER 554 FIGURE 34.—Central rad
Page 101 and 102: NUMBER 554 FIGURE 38.—Central rad
Page 107 and 108: NUMBER 554 FIGURE 44.—Penes of we
Page 115 and 116: NUMBER 554 FIGURE 52.—Penes of ea
Page 117 and 118: Abbott, R.T. 1948. A Potential Snai
Page 119 and 120: NUMBER 554 Society of Washington, 1
Page 121: NUMBER 554 115 gyrus, Gabbia, Hydro
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A Review of the North American Freshwater Snail Genus Pyrgulopsis

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