Freshwater Copepoda (crustacea) from the Chihuahuan Desert with comments on biogeography.
Abstract: The zooplankton fauna of freshwater systems of the Chihuahuan Desert was surveyed 2005-2008. Samples were obtained from habitats including natural lakes, ephemeral waters, springs, and reservoirs. Copepods were sorted and identified to species. Including previous surveys, a total of 38 species occur in the Chihuahuan Desert, the checklist includes 6 calanoids (1 Temoridae, 5 Diaptomidae), 24 cyclopoids (12 Eucyclopinae, 12 Cyclopinae), and 8 harpacticoids. These species were assigned to different biogeographic categories (i.e., cosmopolitan, Nearctic, Neotropical, potential endemics). Results indicated that the Chihuahuan Desert, a transitional biogeographic region in Mexico with both Nearctic and Neotropical influences, represents an interesting geographic and environmental realm characterized by a balanced mixture of elements from different origins and affinities. There is a relatively high proportion (23%) of species that probably are endemic.

La fauna de zooplancton de los sistemas de aguas dulces del desierto de Chihuahua fue muestreada en 2005-2008. Las muestras se obtuvieron de habitats que incluyeron lagos naturales, charcas temporales, manantiales y represas. Se separaron los copepodos y se identificaron a nivel de especie. Incluyendo muestreos previos, un total de 38 especies ocurre en el desierto de Chihuahua, la lista incluye 6 calanoides (1 Temoridae, 5 Diaptomidae), 24 ciclopoides (12 Eucyclopinae, 12 Cyclopinae) y 8 harpacticoides. Estas especies fueron asignadas a diferentes categor?as biogeograficas (por ejemplo, cosmopolita, Neartica, Neotropical, endemica potencial). Los resultados indicaron que el desierto Chihuahuense, una region biogeografica transicional en Mexico, con influencias Nearticas y Neotropicales, representa una interesante entidad geografica y ambiental caracterizada por una mezcla balanceada de elementos de diferentes or?genes y afinidades. Hay una proporcion relativamente alta (23%) de especies que probablemente son endemicas.
Article Type: Report
Subject: Copepoda (Identification and classification)
Copepods (Identification and classification)
Authors: Suarez-Morales, Eduardo
Gutierrez-Aguirre, Martha A.
Walsh, Elizabeth J.
Pub Date: 12/01/2010
Publication: Name: Southwestern Naturalist Publisher: Southwestern Association of Naturalists Audience: Academic Format: Magazine/Journal Subject: Biological sciences Copyright: COPYRIGHT 2010 Southwestern Association of Naturalists ISSN: 0038-4909
Issue: Date: Dec, 2010 Source Volume: 55 Source Issue: 4
Geographic: Geographic Scope: Mexico Geographic Name: Chihuahuan Desert; Chihuahuan Desert Geographic Code: 1MEX Mexico
Accession Number: 246536460
Full Text: A series of surveys of the zooplankton fauna of freshwater systems of northern Mexico and the southwestern United States was developed over several years. These studies included the Chihuahuan Desert; this area is of high priority among hydrological basins of northern Mexico and southwestern Texas (Dinerstein et al., 2000). The Chihuahuan Desert comprises many arid and semi-arid areas with different hydrologic basins and a variety of aquatic habitats; some of these areas are protected to varying degrees (Gomez-Pompa and Dirzo, 1995). We obtained samples of plankton from limnetic and nearshore zones in aquatic habitats including springs, natural lakes, ephemeral pools, and reservoirs. Calanoid, cyclopoid, and harpacticoid copepods were studied taxonomically. The works by Zamudio-Valdez and Reid (1990) and Zamudio-Valdez (1991) in Cuatro Cienegas, Coahuila, which is part ofthe Chihuahuan Desert, were the first surveys of Copepoda in this system. Recently, Suarez-Morales and Walsh (2009) described two new, presumedly endemic, species of Eucylops from this arid region. Herein, we present a complete inventory and perform a general analysis of taxonomic composition and biogeographic affinities of the copepod fauna in the Chihuahuan Desert and compare it with adjacent areas of North America.

MATERIALS AND METHODS--Samples of zooplankton were obtained from freshwater habitats in the Chihuahuan Desert. Biological samples were obtained from springs, ponds, natural lakes, reservoirs, and ephemeral waters during 2005-2008. Samples were taken throughout the year without a planned component for looking at seasonal efforts; 26% of samples were collected in winter, 16% in spring, 48% in summer, and 10% in autumn. Zooplankton was collected by using hand-towed plankton nets (0.64-mm mesh) at both open water (limnetic) and nearshore (littoral) zones of water bodies. Overall, we sampled 134 sites, 103 (77%) were natural and 31 (23%) artificial. Samples were fixed and preserved in 70% ethanol or 4% formalin for long-term storage. Copepods were sorted from the original samples and maintained in 70% ethanol with a drop of glycerine. Specimens were dissected and examined following techniques described by Williamson and Reid (2001) and Reid (2003). Dissected specimens were mounted in semi-permanent slides with glycerine sealed with Entellan (Merck, Darmstadt, Germany) or nail varnish. Specimens were deposited in the collection of zooplankton at El Colegio de la Frontera Sur in Chetumal, Quintana Roo, Mexico.

RESULTS--A total of 32 species of Copepoda was recorded from samples obtained during this survey and 6 more were included from previous surveys. The complete list includes 6 calanoids (1 Temoridae, 5 Diaptomidae), 24 cyclopoids (12 Eucyclopinae, 12 Cyclopinae), and 8 harpacticoids belonging to 5 families (Table 1). Taxonomic arrangement and nomenclature follows Dussart and Defaye (2002, 2006) for Calanoida

and Cyclopoida, and Bodin (1997) for Harpacticoida. These species were assigned to different biogeographic categories (i.e., cosmopolitan, Nearctic, Neotropical, potential endemics; Table 1, Fig. 1).

DISCUSSION--Diaptomid copepods of the Chihuahuan Desert are represented by a mixture of Nearctic and Neotropical forms, showing the expected dominance of Nearctic species in this region of Mexico (Suarez-Morales et al., 2005). Mastigodiaptomus albuquerquensis is the only true Neotropical calanoid occurring in this area; it is a widespread form that has been reported from other areas of Mexico and Central America (Yeatman, 1959; Suarez-Morales and Reid, 1998). It represents the south-to-north radiation of Diaptomidae in the Americas (Suarez-Morales, 2003). Most species of the genus have been recorded in Mexico and Central America (Suarez-Morales and Ellas-Gutierrez, 2000). Specimens of M. albuquerquensis from different areas of Mexico show subtle morphological variations that suggest this taxon as a complex of cryptic species that should be studied further (E. Suarez-Morales, personal observation). Variations in morphology and ecology have been delineated in its congener M. nesus (Cervantes-Martinez et al., 2005), even among sites in geographic proximity.

Leptodiaptomus is a Nearctic genus that radiated southward (Suarez-Morales, 2003); several species have been recorded in Mexico including L. siciloides and L. novamexicanus (Suarez-Morales and Reid, 1998; Suarez-Morales et al., 2005). Leptodiaptomus siciloides has closely related forms (Suarez-Morales et al., 2000) and its variability in North America should be explored further, particularly in isolated or semi-isolated environments like the Chihuahuan Desert.

Both M. albuquerquensis and L. novamexicanus were described originally from adjacent arid or semi-arid areas of New Mexico in the southwestern United States (Herrick, 1895). These species need to be redescribed based on specimens from the type locality to allow comparisons and define the taxonomical status of morphologically similar forms. Unfortunately, type localities of these species are uncertain and the general areas are now deeply transformed (E. J. Walsh, personal observation); hence, neotypes should be designated from adjacent localities.

Presence of the temorid Eurytemora affinis in the Chihuahuan Desert is interesting. This widespread species was presumed to be confined to coastal and estuarine habitats, but it has spread rapidly in the largest riverine basins of North America (Lee and Frost, 2002; Lee and Petersen, 2003). This copepod was collected from two localities in the Chihuahuan Desert as described by Suarez-Morales et al. (2008), who claimed these as the most inland records of this copepod in North America, almost 850 km from the coastline. This also is intriguing because of the hydrologic isolation of these sites, with no evident connection to rivers; a possible explanation of the occurrence of this species in the area includes passive transportation by flows of subterranean water or migrant birds (Reid and Reed, 1994).

As the most diverse group of free-living copepods in freshwaters, cyclopoids were represented by 24 species in the Chihuahuan Desert (Table 1). This number of species is similar to that in adjacent areas of North America: 30 in central Mexico (Suarez-Morales and Reid, 1998), 39 in the Yucatan Peninsula (Suarez-Morales and Reid, 2003), 18 in Tabasco, southeastern Mexico (Gutierrez-Aguirre and Suarez-Morales, 2001), and 41 in Florida (Bruno et al., 2005). In Florida, most cyclopoids are generalists and diversity is not particularly high; however, in the Yucatan Peninsula, local speciation seems to be prevalent (Fiers et al., 1996). Freshwater habitats in the Chihuahuan Desert also harbor many generalist, cosmopolitan cyclopoids, but there is an important proportion of endemic forms that have undergone isolation in geographically close areas, similar to those described in the Yucatan Peninsula (Suarez-Morales et al., 2004).

Among Cyclopidae, cosmopolitan forms were dominant (10 species), followed by Neotropical (5), Nearctic (5), and endemic (4) elements (Table 1). The Neotropical influence is stronger in this group than in the calanoids and it is marked by species of Eucyclops. This genus seems well represented in this arid ecosystem; isolation of some sampled sites appears to favor local speciation in this genus, particularly in forms related to the presumably cosmopolitan E. pectinifer. This species is assumed to represent a diversified complex whose ancestors radiated during post-glacial periods (Alekseev et al., 2006). American records of E. pectinifer (as E. agilis or E. serrulatus) might refer to cryptic species with restricted distributional ranges (Alekseev et al., 2006; Dussart and Defaye, 2006). In the Chihuahuan Desert, the two new species of Eucylops described by Suarez-Morales and Walsh (2009) show subtle but consistent differences with respect to their closest congeners, including E. pseudoensifer and, particularly, E. pectinifer.

The Acanthocyclops robustus-vernalis species complex, including the strict forms, is distributed in different areas of central and northern Mexico (Suarez-Morales and Reid, 1998); recent surveys in this Mexican region showed that there are cryptic species with restricted distributional ranges related to this complex (Mercado-Salas et al., 2006) and to other Nearctic forms of Acanthocyclops (Mercado-Salas et al., 2009). The robustus-vernalis group also has been successful in radiating in tropical areas, where some apparently endemic forms have been described (Reid and Suarez-Morales, 1999; Fiers et al., 2000). Hence, it is expected that the Chihuahuan Desert harbors a greater diversity of endemic copepods than reported here.

Harpacticoids are represented by eight species of which L. stromatolicolus is the only one currently recognized as endemic to the Chihuahuan Desert (Zamudio and Reid, 1990). There are other species that potentially are endemic; these include forms related to the widespread N. lacustris; subtle differences of specimens from Cuatro Cieinegas with respect to the strict form were outlined by Zamudio-Valdez (1991). Other species, such as O. mohammed and C. deitersi and even N. lacustris commonly are in estuarine and brackish water or in transitional environments; their occurrence in this arid region is interesting because they can persist in extreme conditions and are strong colonizers of epigean aquatic environments (Fleeger and Chandler, 1983; Atilla, 2001). There also is fossil evidence suggesting that harpacticoid copepods of the genus Cletocamptus and also some cyclopoid-like forms were present in the region during the Miocene (Palmer, 1960).

Of species collected so far in the Chihuahuan Desert, two cyclopoids and one harpacticoid are likely to be endemic forms (Zamudio and Reid, 1990; Suarez-Morales and Walsh, 2009) and it is expected that other endemic species will be discovered with further research. In addition to its geological age, the Chihuahuan Desert was influenced by the last glaciation (Metcalfe et al., 2002) with alternate wet conditions and deeper, more extensive bodies of water followed by periods of desiccation and deflation. These processes likely resulted in local successive events of isolation in the region. Populations of copepods, particularly cyclopoids, are capable of colonization and dispersal during post-glacial periods in North America (Lewis and Reid, 2007); these forms could have been stranded in these isolated aquatic systems and undergone speciation. The high degree of endemism observed in other invertebrates in the Chihuahuan Desert appears to be extended to the Copepoda.

Overall, the copepod community in the Chihuahuan Desert is a balanced mixture of Neotropical, Nearctic, cosmopolitan, and potentially endemic forms. This likely is explained by geographic location of the Chihuahuan Desert, a latitudinal fringe that represents transition between Nearctic and Neotropical regions. Most records, including those of presumedly endemic forms are from natural environments (Suarez-Morales and Walsh, 2009); thus, suggesting that these systems should be surveyed in more detail. Despite the fact that samples were taken without a pre-established seasonal pattern, almost one-half of these were collected during summer, when rains allow dormant forms to reappear in ephemeral systems. Overall, this year-round sampling provided a more complete coverage of the zooplankton fauna.

In their analysis of the Mexican freshwater copepod fauna, Suarez-Morales and Reid (1998) noted that states and subregions with arid climates may reveal a surprising diversity and provide models for studying dispersal mechanisms and island biogeography. This analysis is in progress, but it is safe to advance that the Chihuahuan Desert is an environment in which geological and environmental isolation of aquatic habitats represents a key factor in favoring local speciation.

This contribution is part of a project funded by National Science Foundation grant DEB-0516032 (E. Walsh). Opinions, results, conclusions, or recommendations expressed herein are those of the authors and do not necessarily reflect views of the National Science Foundation. We thank members of the laboratories of E. J. Walsh, R. Wallace (Ripon College), M. Silva-Briano (Universidad Autonoma de Aguascalientes), and R. Rico-Martinez (Universidad Autonoma de Aguascalientes) for help in collecting samples. Mexican samples were collected under permit to M. Silva-Briano and those from protected areas in the United States were collected under permits to E. J. Walsh. A database that includes the present records was supported by the Comision Nacional para el Conocimiento y Uso de la Biodiversidad (Mexico) through project GT-034.

Submitted 3 April 2009. Accepted 20 March 2010.

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Associate Editor was Jerry L. Cook.

EDUARDO SUAREZ-MORALES, * MARTHA A. GUTIERREZ-AGUIRRE, AND ELIZABETH J. WALSH El Colegio de la Frontera Sur, Unidad Chetumal, Avenida Centenario Km 5.5, Chetumal, Quintana Roo 77014, Mexico (ES-M) Universidad de Quintana Roo, Campus Cozumel, Cozumel, Quintana Roo, Mexico (MAGA) Department of Biological Sciences, University of Texas at El Paso, El Paso, TX 79968 (EJW)

* Correspondent: esuarez@ecosur.mx
TABLE 1--Checklist of free-living copepods (Crustacea) recorded in
freshwater habitats of the Chihuahuan Desert.

                                                    Zamudio-
                                                   Valdeiz and
Taxon                                              Reid, 1990

Order Calanoida
  Family Diaptomidae
    Subfamily Diaptominae
      Leptodiaptomus connexus (Light, 1938)
      Leptodiaptomus novamexicanus (Herrick,
        1895)
      Leptodiaptomus siciloides
        (Lilljeborg, 1889)
      Aglaodiaptomus clavipes (Schacht, 1897)
      Mastigodiaptomus albuquerquensis
        (Herrick, 1895)
  Family Temoridae
      Eurytemora affinis (Poppe, 1880)

Order Cyclopoida
    Subfamily Eucyclopinae
       Ectocyclops phaleratus (Koch, 1838)
       Eucyclops pectinifer (Crangin, 1883)
       Eucyclops cuatrocienegas Suarez-Morales
         and Walsh, 2009
       Eucyclops chihuahuensis Suarez-Morales
         and Walsh, 2009
       Eucyclops bondi Kiefer, 1934
       Eucyclops prionophorus Kiefer, 1931
       Eucyclops pseudoensifer Dussart, 1984
       Macrocyclops albidus albidus
         (Jurine, 1820)
       Paracyclops chiltoni (Thomson, 1882)
       Paracyclops poppei (Rehberg, 1880)
       Tropocyclops cf. extensus Kiefer, 1931

       Tropocyclops prasinus sensu stricto
         (Fischer, 1860)
  Subfamily Cyclopinae
       Acanthocyclops robustus (Sars, 1863)
       Acanthocyclops vernalis (Fischer, 1853)
       Acanthocyclops cf. pennaki Reid, 1992
       Apocyclops dimorphus Kiefer, 1931
       Diacyclops A

       Microcyclops ceibaensis (Marsh, 1919)
       Microcyclops varicans (G. O. Sars, 1863)
       Microcyclops rubellus (Lilljeborg, 1901)
       Mesocyclops edax Forbes, 1891
       Mesocyclops longisetus (Thiei baud, 1893)
       Mesocyclops reidae Petkovsky, 1986
       Thermocyclops tenuis (Marsh, 1909)

Order Harpacticoida
  Family Ameiridae
       Nitokra cf. lacustris (Schmankewitsch,
         1875)

       Nitokra cf. spinipes Boeck, 1865

  Family Darcythompsonidae
       Leptocaris stromatolicolus Zamudio and           X
         Reid, 1990
  Family Canthocamptidae
       Attheyella sp.

       Cletocamptus albuquerquensis (Herrick,
         1895)
       Cletocamptus deitersi Richard, 1897
  Family Laophontidae
       Onychocamptus mohammed (Blanch and
         Rich., 1897)
  Family Miraciidae
       Schizopera sp.

                                                     Zamudio-
Taxon                                              Valdeiz, 1991

Order Calanoida
  Family Diaptomidae
    Subfamily Diaptominae
      Leptodiaptomus connexus (Light, 1938)              X
      Leptodiaptomus novamexicanus (Herrick,
        1895)
      Leptodiaptomus siciloides
        (Lilljeborg, 1889)
      Aglaodiaptomus clavipes (Schacht, 1897)
      Mastigodiaptomus albuquerquensis
        (Herrick, 1895)
  Family Temoridae
      Eurytemora affinis (Poppe, 1880)

Order Cyclopoida
    Subfamily Eucyclopinae
       Ectocyclops phaleratus (Koch, 1838)               X
       Eucyclops pectinifer (Crangin, 1883)            X (a)
       Eucyclops cuatrocienegas Suarez-Morales
         and Walsh, 2009
       Eucyclops chihuahuensis Suarez-Morales
         and Walsh, 2009
       Eucyclops bondi Kiefer, 1934
       Eucyclops prionophorus Kiefer, 1931
       Eucyclops pseudoensifer Dussart, 1984
       Macrocyclops albidus albidus                      X
         (Jurine, 1820)
       Paracyclops chiltoni (Thomson, 1882)              X
       Paracyclops poppei (Rehberg, 1880)                X
       Tropocyclops cf. extensus Kiefer, 1931

       Tropocyclops prasinus sensu stricto
         (Fischer, 1860)
  Subfamily Cyclopinae
       Acanthocyclops robustus (Sars, 1863)
       Acanthocyclops vernalis (Fischer, 1853)           X
       Acanthocyclops cf. pennaki Reid, 1992
       Apocyclops dimorphus Kiefer, 1931                 X
       Diacyclops A

       Microcyclops ceibaensis (Marsh, 1919)             X
       Microcyclops varicans (G. O. Sars, 1863)
       Microcyclops rubellus (Lilljeborg, 1901)          X
       Mesocyclops edax Forbes, 1891
       Mesocyclops longisetus (Thiei baud, 1893)         X
       Mesocyclops reidae Petkovsky, 1986
       Thermocyclops tenuis (Marsh, 1909)

Order Harpacticoida
  Family Ameiridae
       Nitokra cf. lacustris (Schmankewitsch,          X (b)
         1875)

       Nitokra cf. spinipes Boeck, 1865                 X (c)

  Family Darcythompsonidae
       Leptocaris stromatolicolus Zamudio and            X
         Reid, 1990
  Family Canthocamptidae
       Attheyella sp.

       Cletocamptus albuquerquensis (Herrick,            X
         1895)
       Cletocamptus deitersi Richard, 1897               X
  Family Laophontidae
       Onychocamptus mohammed (Blanch and                X
         Rich., 1897)
  Family Miraciidae
       Schizopera sp.

                                                    This
Taxon                                              survey

Order Calanoida
  Family Diaptomidae
    Subfamily Diaptominae
      Leptodiaptomus connexus (Light, 1938)
      Leptodiaptomus novamexicanus (Herrick,         X
        1895)
      Leptodiaptomus siciloides                      X
        (Lilljeborg, 1889)
      Aglaodiaptomus clavipes (Schacht, 1897)        X
      Mastigodiaptomus albuquerquensis               X
        (Herrick, 1895)
  Family Temoridae
      Eurytemora affinis (Poppe, 1880)               X

Order Cyclopoida
    Subfamily Eucyclopinae
       Ectocyclops phaleratus (Koch, 1838)           X
       Eucyclops pectinifer (Crangin, 1883)          X
       Eucyclops cuatrocienegas Suarez-Morales       X
         and Walsh, 2009
       Eucyclops chihuahuensis Suarez-Morales        X
         and Walsh, 2009
       Eucyclops bondi Kiefer, 1934                  X
       Eucyclops prionophorus Kiefer, 1931           X
       Eucyclops pseudoensifer Dussart, 1984         X
       Macrocyclops albidus albidus                  X
         (Jurine, 1820)
       Paracyclops chiltoni (Thomson, 1882)          X
       Paracyclops poppei (Rehberg, 1880)            X
       Tropocyclops cf. extensus Kiefer, 1931        X

       Tropocyclops prasinus sensu stricto           X
         (Fischer, 1860)
  Subfamily Cyclopinae
       Acanthocyclops robustus (Sars, 1863)          X
       Acanthocyclops vernalis (Fischer, 1853)
       Acanthocyclops cf. pennaki Reid, 1992         X
       Apocyclops dimorphus Kiefer, 1931             X
       Diacyclops A                                  X

       Microcyclops ceibaensis (Marsh, 1919)
       Microcyclops varicans (G. O. Sars, 1863)      X
       Microcyclops rubellus (Lilljeborg, 1901)      X
       Mesocyclops edax Forbes, 1891                 X
       Mesocyclops longisetus (Thiei baud, 1893)
       Mesocyclops reidae Petkovsky, 1986            X
       Thermocyclops tenuis (Marsh, 1909)            X

Order Harpacticoida
  Family Ameiridae
       Nitokra cf. lacustris (Schmankewitsch,        X
         1875)

       Nitokra cf. spinipes Boeck, 1865            X

  Family Darcythompsonidae
       Leptocaris stromatolicolus Zamudio and      X
         Reid, 1990
  Family Canthocamptidae
       Attheyella sp.                              X

       Cletocamptus albuquerquensis (Herrick,
         1895)
       Cletocamptus deitersi Richard, 1897         X
  Family Laophontidae
       Onychocamptus mohammed (Blanch and          X
         Rich., 1897)
  Family Miraciidae
       Schizopera sp.                              X

                                                   Biogeographic
Taxon                                              category

Order Calanoida
  Family Diaptomidae
    Subfamily Diaptominae
      Leptodiaptomus connexus (Light, 1938)        Nearctic
      Leptodiaptomus novamexicanus (Herrick,       Nearctic
        1895)
      Leptodiaptomus siciloides                    Nearctic
        (Lilljeborg, 1889)
      Aglaodiaptomus clavipes (Schacht, 1897)      Nearctic
      Mastigodiaptomus albuquerquensis             Neotropical
        (Herrick, 1895)
  Family Temoridae
      Eurytemora affinis (Poppe, 1880)             Cosmopolitan

Order Cyclopoida
    Subfamily Eucyclopinae
       Ectocyclops phaleratus (Koch, 1838)         Cosmopolitan
       Eucyclops pectinifer (Crangin, 1883)        Cosmopolitan
       Eucyclops cuatrocienegas Suarez-Morales     Endemic or
         and Walsh, 2009                           potentially endemic
       Eucyclops chihuahuensis Suarez-Morales      Endemic or
         and Walsh, 2009                           potentially endemic
       Eucyclops bondi Kiefer, 1934                Neotropical
       Eucyclops prionophorus Kiefer, 1931         Neotropical
       Eucyclops pseudoensifer Dussart, 1984       Neotropical
       Macrocyclops albidus albidus                Cosmopolitan
         (Jurine, 1820)
       Paracyclops chiltoni (Thomson, 1882)        Cosmopolitan
       Paracyclops poppei (Rehberg, 1880)          Cosmopolitan
       Tropocyclops cf. extensus Kiefer, 1931      Endemic or
                                                   potentially endemic
       Tropocyclops prasinus sensu stricto         Cosmopolitan
         (Fischer, 1860)
  Subfamily Cyclopinae
       Acanthocyclops robustus (Sars, 1863)        Cosmopolitan
       Acanthocyclops vernalis (Fischer, 1853)     Cosmopolitan
       Acanthocyclops cf. pennaki Reid, 1992       Nearctic
       Apocyclops dimorphus Kiefer, 1931           Nearctic
       Diacyclops A                                Endemic or
                                                   potentially
                                                   endemic?
       Microcyclops ceibaensis (Marsh, 1919)       Neotropical
       Microcyclops varicans (G. O. Sars, 1863)    Cosmopolitan
       Microcyclops rubellus (Lilljeborg, 1901)    Cosmopolitan
       Mesocyclops edax Forbes, 1891               Nearctic
       Mesocyclops longisetus (Thiei baud, 1893)   Neotropical
       Mesocyclops reidae Petkovsky, 1986          Neotropical
       Thermocyclops tenuis (Marsh, 1909)          Neotropical

Order Harpacticoida
  Family Ameiridae
       Nitokra cf. lacustris (Schmankewitsch,      Endemic or
         1875)                                     potentially endemic

       Nitokra cf. spinipes Boeck, 1865            Endemic or
                                                   potentially endemic
  Family Darcythompsonidae
       Leptocaris stromatolicolus Zamudio and      Endemic or
         Reid, 1990                                potentially endemic
  Family Canthocamptidae
       Attheyella sp.                              Endemic or
                                                   potentially endemic
       Cletocamptus albuquerquensis (Herrick,      Neotropical
         1895)
       Cletocamptus deitersi Richard, 1897         Cosmopolitan
  Family Laophontidae
       Onychocamptus mohammed (Blanch and          Cosmopolitan
         Rich., 1897)
  Family Miraciidae
       Schizopera sp.                              Endemic or
                                                   potentially endemic

(a) Possibly is Eucyclops cf. pectinifer A (E. Suarez-Morales, personal
observation).

(b) Possibly undescribed, reported as N. lacustris by Zamudio-Valdez
(1991).

(c) Possibly undescribed, reported as N. spinipes by Zamudio-Valdez
(1991).

FIG. 1--General overview of biogeographic affinities
of copepods recorded in the Chihuahuan Desert
during this survey. Number of species in each of the
four categories is indicated next to each slice of
the graphic.

BIOGEOGRAPHIC OVERVIEW

Neotropical     9
Cosmopolitan   13
Nearctic        9
"Endemics"      7

Note: Table made from pie chart.
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