The Arachnid Order Solifugae

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PHYLOGENY/TAXONOMY

Phylogeny of the Solifugae
Keys to FamiliesFamily Ammotrechidae
Family Ceromidae
Family Daesiidae
Family Eremobatidae
 


 

 

 

 

 

 

 

Family Galeodidae
Family Gylippidae
Family Hexisopodidae
Family Karschiidae
Family Melanoblossidae
Family Mummuciidae
Family RhagodidaeFamily Solpugidae
Catalog of the Solifugae

Genus Eremobates Banks 1900



Female Eremobates sp., palpisetulosus group from California, USA.
Photo by Warren Savary.

Genus Eremobates Banks 1900

Datames ►Simon, 1879a: 133-135; Simon, 1879c: 78 [junior homonym of Datames Stål, 1875 (Insecta: Orthoptera)].
Eremobates
►Banks 1900: 426; ►Kraepelin 1901a: 117; ►Pocock 1902: 59; Comstock 1913: 36; Pratt 1927: 408; ►Roewer 1934: 555, 556, fig. 320 a, h; Pratt 1935: 477; ►Muma 1951: 41, 51; ►Muma 1970a: 10; ►Muma 1976: 7, 15; ►Muma 1989: 3, 6; Harvey 2001a: 198-199 [replacement name for Datames Simon, 1879a]; ►Brookhart and Cushing 2004: 288; ►Brookhart and Brookhart 2006: 301.
Eremoperna
►Roewer 1934: 555-557, figs. 320 b, i (synonymized by Muma, 1951: 51).
Eremospina
►Roewer 1934: 555, 556, 565, figs. 320 c, l  (synonymized through transfer of type species to Eremobates by Muma, 1970a: 29).
Eremognatha
►Roewer 1934: 555, 556, 566, figs. 320 d, l  (synonymized by Muma, 1951: 51).
Eremoseta
Roewer ►1934: 555, 556, 569, figs. 320  e, l (synonymized by Muma, 1970a: 21).
Eremostata
Roewer, ►1934: 555, 556, 571-572, figs. 320  (synonymized by Muma, 1951: 51).

Type species:

of Datames and Eremobates:  Gluvia cinerascens C.L. Koch 1842, by subsequent designation of Pocock (1902c): 59 [see Harvey 2001a: 198-199] [see Eremobates cinerascens (C.L.Koch)].
of Eremoperna:  Datames affinis Kraepelin 1899b, by original designation [see Eremobates affinis (Kraepelin)].
of Eremospina:  Gluvia tolteca Pocock 1895a, by original designation [see Eremobates tolteca (Pocock)].
of Eremognatha:  Datames tuberculatus Kraepelin 1899b, by original designation [see Eremobates tuberculatus (Kraepelin)]. 
of Eremoseta:  Eremoseta titschacki Roewer 1934, by original designation [see Eremobates titschacki (Roewer)]. 
of EremostataGluvia cinerascens C.L. Koch 1842, by original designation [see Eremobates cinerascens (C.L. Koch)].

Diagnosis:  S

Original description:
Simon, 1879a: 133-135 (as Datames):

"Mamelon oculaire garni en dessus el en avant de crins nombreux, irréguliers, non soulevés. - Partie céphalique peu longue, obtusément trapézoide, en ligne droite ou un peu arquée en avant, en ligne presque droite en arriere, a cotes assez fortement obliques et convergeant en arrière, sans strie longitudinale. - Chélicères de la femelle à crochet fixe en dessus, plus court que la  tige, arqué et pourvu d'une rangée de dente inégales disposèes comme chez let Gluvia, seulement les deux dents principales toujours séparées par deux denticules; chez le màle, le crochet fixe ausel long que la tige, brusquement rétréci dès la base, droit, dépourvu de dents et canaliculé en dessous à la base; dans les deux sexes, le fond da la pince pourvu de deux rangées écartées de dents obtuses; les deux dents principales du crochet mobile séparées par deux denticules, - Pas de peignés stigmatatiques.  - Patte-machiore à tarse oourt soudé au métatarse et immobile. - Pattes de la première paire à tarse long droit. gréle, de moitié ou d'un tiers plus court que le métatarse. dépourvu da griffes dans les deux sexes. - Griffes longues, gréles, g1abres, plus courtes aux paires intermédiaires qu'à la quatrième, - Hanches de la patte-máchoire et de la première paire réunies, presque aussi larges que longues.
Ce genre est exclusivement propre à l'Amérique; il renferme toutes les espèces anciennement décrites par T. Say et plus récemment par C. Koch, sous le nom générique de Gluvia. Les deux groupes établis par C. Koch (in Archiv. f. Naturg.) reposent sur un caractère sexuel, aussi les espèces du second ne sont-elles pour la plupart que les femelles de celles du premier.
Le genre Datames a deux centres principaux: le premier aux États-Unis, dans les montagnes Rocheuses et les régions du Pacifique: en Colorado, Nevada. Utah et Californie; il n'a aucun représentant dans les réglone de l'Atlantique, si ce n'est dans Texas. où a élé signalée une espèce mexicaine (geniculatus). Il est à. remarquer que toutes les spèces des Étals-Unis se reconnaissent de suite en ce que, chez les femelles, les deux petites denticulations intermédiaires des chélicères sont parfaitement isolées. - Le second centre est au Mexique, le genre Datames y est représenté par des espéces généralement plus grandes, dont les deux petites denticulations sont toujours implantées sur la  antérieure de la seconde dent principale.  C. Koch a aussi
décrit deux Datames de Colombie dont la provenance est pent--étre erronée, car, dans toute l'Amérique du Sud et les Antilles, les Datames paraissent remplacés par les Cleobis, et les Mummucia.
Les caract
ères propres aux deux sexes, qui séparent les Datames des Gluvia, sont assez secondaires, mais d'une constance absolue: chez les Datames, les deux dents principales des crochete des chélicères sont séparees par deux denticules, tandis qu'll n'y, a qu'un seul chez les Gluvia; le métatarse de la troisième paire porte en dessus une ligne de cinq fortes épines ches les Datames, tandis que celui des Gluvia n'en a que trois; mais les Datames s'éloignent de tous les Galeodidae connus par l'éntrange conformation des chélicères du male, dont le crochet fixe est inerme et droit, et dont le flagellum est remplacé par un fort faisceau de crins; l'abdomen est dépourvu des épines bacilliformes si caractéristiques chez les Gluvia."

Subsequent accounts:
Banks, 1900: 426:

"Our genera and species may be tabulated as below.

1. Anterior margin of the cephalothorax rounded, sIoping each side............... (Ammotrecha) 9 l
   
Anterior margin truncate...............................................................................................(Eremobates) 22

                                                                            ***[
key continued]***
                                  2
Eremobates, n.n. for Datames Simon (1879). not Stal (1875)."

Kraepelin 1901a: 117:

"Mit den Merkmalen der Unterfamilie.
18 sichere und 3 unsichere Arten.
Bei der grossen Verschiedenheit der ♂ und ♀ und bei der Spärlichkeit des in den Sammlungen vorliegenden Materials ist erst für veihältnismässig wenige Arten die Zusammengehörigkeit von ♂ und ♀ sicher festgestellt. Es sind daher zahlreiche ♂ und ♀ gesondert beschrieben und benannt, die sich voraussichtlich später als zusammengehörig erweisen werden. Da überdies die ♀ der verschiedenen Arten, wie auch sonst bei den Solifugen, eine weitgehende Übereinstimmung zeigen, so kann der nachfolgende Bestimmungsschlüssel nur als erster Versuch gelten, die Hauptformen von einander abzugliedern."

Pocock, 1902: 59:

"Leg of first pair without claw. Tarsus of legs furnished beneath with distinct pairs of spines; tibia of second and third legs furnished above with five strong spines arranged in a single series, and three or four pairs of weaker spines below. Mandibles with only feebly developed stridulating-ridges on the anterior portion of the smooth area on the inner side of their appendages; without any flagellum, and with the upper jaw untoothed in the male.
Type Gluvia cinerascens, C. L. Koch.

Distribution.
Southern States of North America and Mexi
co."

Roewer, 1934: 555, 556, fig. 320 a, h (as Eremobates Banks):

Genotypus: E. pallipes (Say).
Eremobatinae, deren 2. und 3. Tarsus ventral mit jeweils 2.2 Dornen und deren 4. Tarsus ventral mit 2.2/0/2 Dornen bewehrt ist (Abb. 320, a, h). 
2 Arten, von der eine nur als ♂, die andere nur als ♀ bekannt ist.
Eremobates pallipes (Say 1823), Eremobates durangonus Roewer 1934


Abb. 320a. Ventrale Bedornung des 2.-oder 3. Tarsus von Eremobates.
Abb. 320h. Ventrale Bedornung des 4. Tarsus von Eremobates.
Redrawn from Roewer 1934.

Roewer, 1934: 555-557, figs. 320 b, i (as Eremoperna Roewer 1934):

Genotypus: E affinis (Kraepelin).
Eremobatinae, deren 2. und 3. Tarus ventral mit jeweils 4.6 Dornen und deren 4. Tarsus ventral mit 2.2.2/0/2.4 Domen bewehrt ist (Abb. 320, b, i).
4 Arten, von denen eine nur als ♀ bekannt ist.
Datames affinis
Kraepelin 1899, Datames formidabilis Simon 1879, Eremobates azteca Pocock 1902, Eremoperna mormona Roewer 1934


Abb. 320b. Ventrale Bedornung des 2.-oder 3. Tarsus von Eremoperna.
Abb. 320i. Ventrale Bedornung des 4. Tarsus von Eremoperna.
Redrawn from Roewer 1934.

Roewer, 1934: 555, 556, 565, figs. 320 c, l (as Eremospina Roewer 1934):

Genotypus: E. tolteca (Pocock).
Eremobatinae,
deren 2. und 3. Tarsus ventral mit jeweils 1.4.6 Dornen und deren 4. Tarsus ventral mit 2.2.2/0/2.4.6 Dornen bewehrt ist (Abb. 320, c, l).
Nur 1 Art.
Gluvia tolteca
Pocock 1895


Abb. 320c. Ventrale Bedornung des 2.-oder 3. Tarsus von Eremospina.
Abb. 320l. Ventrale Bedornung des 4. Tarsus von Eremospina.
Redrawn from Roewer 1934.

Roewer, 1934: 555, 556, 566, figs. 320 d, l  (as Eremognatha Roewer 1934):

Genotypus: E. tuberculatus (Kraepelin).
Eremobatinae
, deren 2. und 3. Tarsus ventral mit jeweils 2.2.2.4.6 Dornen und deren 4. Tarsus ventral mit 2.2.2/0/2.4.6 Dornen bewehrt ist (Abb. 320, d, l).
3 Arten, von denen die eine nur als ♂ bekannt ist:
Datames tuberculata Kraepelin 1899,
Eremobates marginata Kraepeliu 1911, Eremognatha guenini Roewer 1934.


Abb. 320d. Ventrale Bedornung des 2.-oder 3. Tarsus von Eremognatha.
Abb. 320l. Ventrale Bedornung des 4. Tarsus von Eremognatha.
Redrawn from Roewer 1934.

Roewer, 1934: 555, 556, 569, figs. 320 e, l (as Eremoseta Roewer 1934):

Genotypus : E. titschacki Roewer 1934.
Eremobatinae,
deren 2. und 3. Tarsus ventral mit jeweils 2.2.4.4.6.6.6 Dornen und deren 4. Tarsus ventral mit 2.2.2/0/2.4.6 Dornen bewehrt ist (Abb. 320, e, l).
Nur I Art:
Eremoseta titschacki Roewer 1934.


Abb. 320e. Ventrale Bedornung des 2.-oder 3. Tarsus von Eremoseta.
Abb. 320l. Ventrale Bedornung des 4. Tarsus von Eremoseta.
Redrawn from Roewer 1934.

Roewer, 1934: 555, 556, 571-572, figs. 320 g, o  (as Eremostata Roewer 1934):

Genotypus: E. cinerascens (Koch).
Eremobatinae,
deren 2. und 3. Tarsus ventral mit jeweils 2.4.4.6.6.6 Dornen und deren 4. Tarsus ventral mit 4.2.2.4.2.2.4/2/2.4.6 Dornen bewehrt ist (Abb. 320, g, o). (Die ♀♀
aller bisher bekannten Arten haben keine wahrnehmbaren Ctenidien an den opisthosomalen Sterniten.)
7 Arten, von denen nur eine in beiden Geschlechtern bekannt ist.

Gluvia cinarascens
Koch 1842, Eremostata arizonica Roewer 1934, Datames sulfurea Simon 1879, Datames caroliniana Kraepelin 1899, Datames californica Simon 1879, Eremostata dinamita Roewer 1934, Datames scabra Kraepelin 1899


Abb. 320g. Ventrale Bedornung des 2.-oder 3. Tarsus von Eremostata.
Abb. 320o. Ventrale Bedornung des 4. Tarsus von Eremostata.
Redrawn from Roewer 1934.

 

Muma, 1951: 51:

Small- to moderate-sized Eremobatinae. Fixed finger of males with a mesoventral groove that extends the entire length of the finger. Flagellum complex composed of a dorsal row of simple tubular bristles that are sometimes striate and a ventral row of S-shaped, flattened, plumose bristles that form an arch over the basal third of the mesoventral groove. Apical, plumose bristle of ventral row straight and forming a parallel covering for the apical two-thirds of the mesoventral groove. First post-spiracular abdominal sternite of males with or without ctenidia on its posterior margin. Mesal tooth of movable finger of males and females present or absent. Genital opercula of females variable.

Muma, 1970a: 3, 10:

Leg I with 1 claw; large, robust, short legged species
Palpus with or without spinelike setae; when present. apparently not movable and not on metatarsus and tarsus
Fixed finger elongate and needlelike or stylelike
Mesoventral groove of fixed ftnger extending to base of finger; apical seta of flagellum-complex flattened and plumose

Muma 1986/Muma 1987

None of the new species assigned to this genus significantly extend or alter the presently established generic characters.

Muma 1989: 3, 6

Mostly moderate-sized Eremobatinae. Fixed cheliceral finger of males with mesodorsal or mesoventral groove extending the length of finger. Dorsal setae of flagellum complex striate, ventral setae plumose; apical striate seta not obviously modified but apical plumose seta conspicuously flattened and covering part to nearly all of mesoventral groove. Most females have the opercula broadly triangular in shape but variable along mesal and posterior margins. Females of palpisetulosus, aztecus, and vallis groups have opercula pitted and alate laterally but variable in size and form. Fondal tooth formulae are no longer considered valid for genus or group separation; they are subject to wear, especially by females.

Brookhart and Cushing, 2004: 288:

"Muma (1951) described the genus Eremobates as small to medium sized Eremobatidae with a mesoventral groove that extends the entire length of the male fixed finger.  The flagellum complex is composed of a dorsal row of simple tubular bristles that are sometimes striate and a ventral row of S-shaped, flattened, plumose bristles that form an arch over the basal third of the mesoventral groove. The apical, plumose bristle of the ventral row is straight and forms a parallel covering over the apical two-thirds of the mesoventral groove. The first post-spiracular abdominal sternite of males are with or without ctenidia. Genital operculum of female variable. This description did not change in later works (Muma 1962, 1970, 1989)."

Notes:

Muma (1951), in a key to the genera of the subfamily Eremobatinae, based on males, attributed the following features to Eremobates: Fixed finger of chelicera style-like; Mesoventral groove of fixed finger extending to base of finger, small to moderate-sized species.   Muma (1976) recognized the genus as valid and as a member of the family Eremobatidae, and included the genus in an annotated list of Western Hemisphere Solpugida. He reported at the time that, "Although 58 names have been applied to species of this large genus, only 36 are presently considered valid." Muma (1989) attributed the following features to Eremobates in a key to the subfamilies and genera of the family Eremobatidae: Leg 1 with one claw; chelicerae about twice as long as wide; small to large species; fixed cheliceral finger long, style-like or needle-like; mesoventral groove distinct, extending to base of fixed finger; apical plumose seta of male flagellum complex obviously enlarged and flattened covering part of mesoventral groove; some palpal setae enlarged and spine-like; movable finger dentition normal or increased, moderate-sized to large species.  Brookhart and Brookhart (2006) reproduced the Key to Subfamilies and Genera of Eremobatidae (males only) previously published by Muma (19890, adopting the diagnostic characters used to distinguish Eremobates.  Muma (1982) reported that Eremobates comprises both the greatest number of species and the greatest number of individuals, often representing over 40% of the solpugid population in an area.

Subordinate taxa:

Muma (1951) recognized four species groups within the genus Eremobates (the scaber, palpisetulosus, pallipes, and angustus groups), diagnosed them, and provided a key to them (based on males). Muma (1970) later recognized five species-groups, including all of the above, plus the monobasic aztecus group, which he  erected and diagnosed  for the first time to include Eremobates aztecus .  He also provided an identification key, and listed 13 "nomina dubia" not assigned to species groups:  Eremobates audax (Hirst), Eremobates californicus (Simon), Eremobates carolinianus (Krraepelin), Eremobates caspari (Marx), Eremobates constricta (Putnam), Eremobates dorsalis (Roewer), Eremobates elongatus (C. L. Koch), Eremobates hystrix (Mello-Leitão, Eremobates ingens Mello-Leitao, Eremobates praecox (C. L. Koch), Eremobates subulata (Girard), Eremobates sulfureus (Simon), and Eremobates tolteca (Pocock).  Eremobates praecox (C. L. Koch), originally described as Gluvia praecox C. L. Koch 1842, had previously been treated as Biton praecox (C. L. Koch) by Roewer (1932), and was associated with Eremobates for the first time by Muma (1970).  It was recently listed by Harvey (2003), in his Catalogue of the Smaller Arachnid Orders of the World, as Biton praecox (C.L. Koch). Muma included the same five species groups in his annotated list of Western Hemisphere Solpugida, along with 12 unplaced species (dropping Eremobates praecox (C. L. Koch) from his previous list).  Muma (1986, 1987) added new species to the palpisetulosus and angustus groups, and established a new monbasic species group, the lapazi group, to accommodate the newly named Eremobates lapazi Muma from the cape region of Baja California, Mexico, bringing the total number of species groups to six. Muma (1989) established another monobasic species group, the vallis group, to accommodate a newly described species, Eremobates vallis Muma, from the southwestern United States.  Thus, currently recognized species groups of Eremobates are the scaber, aztecus, palpisetulosus, pallipes, angustus, lapazi, and vallis groups, for which Muma (1989) provided identification keys (one based on males, one on females).  The monophyly of the groups has never been tested, and the relationships between the groups have not been established. Muma (1989) reported that in females of the palpisetulosus, aztecus, and vallis groups, the opercula are pitted and alate laterally, but variable in size and form. The fondal tooth formulae, he noted, are not valid for genus or group separation, as they are subject to wear, especially in females. Brookhart and Brookhart (2006) included six of the seven species groups in their Annotated Checklist of Continental North American Solifugae, omitting Eremobates vallis and the vallis group from the list without explanation.  The current onstituency of the seven species groups is as follows:

angustus group

Eremobates angustus Muma 1951
Eremobates becki Muma 1986
Eremobates cruzi Muma 1951

 

palpisetulosus group

Eremobates affinis (Kraepelin 1899)
Eremobates ajoanus Muma & Brookhart 1988
Eremobates bajadae Muma & Brookhart 1988
Eremobates bajaensis Muma 1986
Eremobates bantai Brookhart 1965
Eremobates bixleri Muma & Brookhart 1988
Eremobates coahuilanus Muma 1986
Eremobates fagei (Roewer 1934)
Eremobates girardi (Putnam 1883)
Eremobates gracilidens Muma 1951
Eremobates guenini (Roewer 1934)
Eremobates hessei (Roewer 1934)
Eremobates hystrix (Mello-Leitao 1942)
Eremobates inyoanus Muma & Brookhart 1988
Eremobates kastoni Muma & Brookhart 1988
Eremobates kiseri Muma & Brookhart 1988
Eremobates kraepelini Muma 1951
Eremobates leechi Muma & Brookhart 1988
Eremobates marathoni Muma 1951
Eremobates nanus Muma 1962
Eremobates nivis Muma & Brookhart 1988
Eremobates nodularis Muma 1951
Eremobates norrisi Muma & Brookhart 1988
Eremobates otavonae Muma & Brookhart 1988
Eremobates pallidus Muma & Brookhart 1988
Eremobates palpisetulosus Fichter 1941
Eremobates papillatus Muma 1970
Eremobates pimanus Muma & Brookhart 1988
Eremobates polhemusi Muma & Brookhart 1988
Eremobates purpusi (Roewer 1934)
Eremobates pyriflora Muma & Brookhart 1988
Eremobates scopulatellus Muma & Brookhart 1988
Eremobates scopulatus Muma 1951
Eremobates spissus Muma & Brookhart 1988
Eremobates tejonus Chamberlin 1925
Eremobates texanus Muma & Brookhart 1988
Eremobates titschacki (Roewer 1934)
Eremobates tuberculatus (Kraepelin 1899)
Eremobates vicinus Muma 1963
Eremobates villosus Muma 1951
Eremobates williamsi Muma & Brookhart 1951

vallis group

Eremobates vallis Muma 1989

 

aztecus group

     Eremobates aztecus Pocock 1902

lapazi group

Eremobates lapazi Muma 1986

 

scaber group

Eremobates actenidia Muma 1989
Eremobates ascopulatus Muma 1951
Eremobates clarus Muma 1989
Eremobates corpink Brookhart & Cushing 2004
Eremobates ctenidiellus Muma 1951
Eremobates hodai Muma 1989
Eremobates icenogelei Brookhart & Cushing 2004
Eremobates legalis Harvey 2002
Eremobates mormonus (Roewer 1934)
Eremobates scaber (Kraepelin 1899)
Eremobates similis Muma 1951
Eremobates socal Brookhart & Cushing 2004
Eremobates zinni Muma 1951

 

pallipes group

Eremobates arizonicus (Roewer 1934)
Eremobates californicus (Simon 1879)
Eremobates chihuaensis Brookhart & Cushing 2002
Eremobates cinerascens (C. L. Koch 1842)
Eremobates constrictus (Putnam 1882)
Eremobates dentilis Brookhart & Muma 1981
Eremobates dilatatus (Putnam 1882)
Eremobates docolora Brookhart & Muma 1981
Eremobates durangonus Roewer 1934
Eremobates formicarius (C. L. Koch 1842)
Eremobates gerbae Brookhart & Cushing 2002
Eremobates lentiginosus (Kraepelin 1899)
Eremobates pallipes (Say 1823)
Eremobates putnami (Banks 1898)
Eremobates simoni Muma 1970
Eremobates suspectus Muma 1951
Eremobates woodruffi Brookhart & Muma 1988

 

Keys:  Muma (1989) offered the following keys to the species groups of Eremobates [illustrations have been added herein to facilitate use of the keys]:

Key to the species groups of Eremobates (based on males) from Muma (1989). 
Figures added (redrawn from Muma 1951, 1970, 1986, 1989).
1 Mesal groove of fixed cheliceral finger mesodorsal in position

aztecus group

Mesal groove of fixed cheliceral finger mesoventral in position

 

 

 

see couplet 4

2 Mesal groove broad both basally and apically, occupying most of finger width



vallis
group

Mesal groove narrow apically, occupying less than half of
finger width

see couplet 3

3 Mesal groove not dilated basally
 

angustus group

Mesal groove distinctly dilated basally


 

see couplet 4

4 Fixed cheliceral finger in dorsal view with a wide notch or off-set

scaber group

Fixed cheliceral finger in dorsal view straight or at most slightly sinuate



                        see couplet 5

5 Fixed finger with an ectodorsal, ridge-like or tooth-like process near its base

palpisetulosus group

Fixed finger constricted near base but without an ectodorsal tooth-like process



 

see couplet 6

6 Anterior tooth of movable cheliceral finger present and normal; small species



pallipes group

Anterior tooth of movable cheliceral finger absent or abortive; large species
 



l



lapazi group


 

Key to the species groups of Eremobates (based on females) from Muma (1989). 
Figures added (redrawn from Muma 1951, 1970, 1986, 1989).
1 Opercula widely separated posteriorly and/or slightly to distinctly lobate at posterio-ectal angles and with a pair of distinct pits about midway along ectal margins

see couplet 2

Opercula not or only moderately separated posteriorly; broadly triangular in shape and without distinct pits midway along ectal margins (except for E. mimbrenus Muma)
 

see couplet 4

2 Posteriomesal notch small and flat, occupying a minor portion of opercular area, pits shallow




vallis
group

Posteriomesal notch wide and long, occupying at least a third of opercular area, pits deep



                          see couplet 3

3 Posteriomesal notch of opercula smoothly flared or arched mesally
 

palpisetulosus group

Posteriomesal notch of opercula sinuate or weakly lobate mesally within the notch

aztecus group

4 Mesal margins of opercula lobate, bilobate or sinuate at or just anterior to posteriomesal notch.

scaber group

Mesal margins of opercula straight or evenly curved to the posteriomesal notch


see couplet 5

5 Mesal margins of opercula divergent both anteriorly and posteriorly
 

angustus group

Mesal margins of opercula parallel anteriorly although variously notched posteriorly

see couplet 6

6 Posterior mesal margins of opercula sinuate; posteriomesal notch narrow and slot-like
 

lapazi group

Posterior mesal margins of opercula straight or evenly curved; posteriomesal notch variable in form
 

pallipes group

   
   

 



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