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Home , Canadian River, Delaware River (Texas), Mimbres River, Pecos River, Phenacobius mirabilis, Rio Bonito (New Mexico)

THE NATIVE FISH FAUNA OF MAJOR DRAINAGES

EAST OF THE CONTINENTAL DIVIDE

IN NEW

A Thesis Presented

to the

Graduate Faculty of Biology

Eastern University

In Partial Fulfillment

of the Requirements

fdr -the7Degree:

Master of Science in Biology

by

Michael D. Hatch

December 1984 TABLE OF CONTENTS

Page Introduction

Study Area

Procedures

Results and Discussion

Summary

Acknowledgements

Literature Cited

Appendices

Abstract INTRODUCTION

r (t. The earliest impression of New Mexico's native fish fauna =Ems during

the 1850's from naturalists attached to various government survey parties.

Without the collections from these and other early surveys, the record of the native fish fauna would be severely deficient because, since that time, some

1 4 native species - or subspecies of fish have become extirpated and the ranges of an additionial 22 native species or subspecies have become severly re- stricted.

Since the late Miocene, physiographical changes of drainages have linked

New Mexico, to varying degrees, with contemporary ichthyofaunal elements or

their progenitors from the , the , the Chihuahuan

Desert, the , the Sonoran and the Great Basin. Immigra-

tion from these areas contributed to the diversity of the state's native

ichthyofauna. Over the millinea, the fate of these fishes waxed and waned in

ell 4, response to the changing physical and _chenaca-l-conditions of the surrounding environment. Ultimately, one of the most diverse fish faunas of any of the

interior southwestern states developed. Fourteen families comprising 67 species of fish are believed to have occupied New Mexico's waters historically, with strikingly different faunas evolving east and west of the Continental

Divide.

The fluvial systems that drain the west slope of the Continental Divide

in New Mexico are of the . The origin and composi-

tion8of the native ichthyofauna in these drainages has received much attention

(Miller 1958, Rinne 1969, Holden 1970, Smith et al. 1983, Minckley et al.

in press). In contrast, only partial coverage has ever been attained of

the origin and composition of the state's native ichthyofauna east of the

Continental Divide.

The aims of this paper are to: (1) present a summary and synthesis of 2c. historical information concerning the distribution of fishes east of the

Continental Divide in New Mexico, (2) determine the major prehistoric events and conditions that governed the origin of the native fish fauna in these drainages, and (3) reconstruct this fauna as it likely existed at the start of the historic era (circa 1550 AD).

STUDY AREA Basin - The Rio Grande originates in the of

southwestern Colorado and flows in a southernly direction through New Mexico

and along the between and Mexico for 1887 miles to the Gulf

of Mexico. The river flows through a deeply incised (approximately 1,000

feet deep) and narrow (ex. the flood plain is typically no wider

than 50 feet) of the Taos Plateau as it enters New Mexico. The river exit.;-

this canyon at Embudo, New Mexico and continues south through a broad valley

that extends to Otowi bridge (just south of Santa Clara ), then

becomes abruptly constricted as it enters another canyon that extends

downstream to Cochiti Pueblo. For the remainder of its course in New Mexico,

the Rio Grande flows through a broad valley of low relief. Stream gradients

of 25 feet per mile are common upstream of Cochiti Pueblo; stream gradients

of 5 feet per mile prevail between Cochiti Pueblo and El Paso.

The Rio Grande upstream of Cochiti Pueblo is a perennial river with

numerous perennial tributaries. Contrasting this, there are no perennial

tributaries of the Rio Grande in New Mexico downstream of Cochiti Pueblo

and much of this portion of the river is dewater6a—by, or as a result of,

nthropogenic events for extensive periodl. However, this situation is

not natural. Prior to settlement of the by Anglo-

Europeans, the Rio Grande was perennial throughout its course in New Mexico.

Even the most arid part of the flood plain was characterized by numerous

marshes, oxbow pools and a fringe-forest of cottonwoods, willows and shrubby

phreatophytes. (Metcalf 1967). Miller (1961) cites the occurrance of

Scaphirhynchus platorynchus taken near Albuquerque in 1873 and the ideff44-44-c4=--

--1:12MPTrf remains of Cycleptus eloncatus from an Indian site west of Santa Fe

as evidence that the upper Rio Grande was once a larger river of more reliable

flow.

After the Civil Vi6r, western migration of Anglo-European settlls was

accelerated and a large number of people settled in the state, especially in the Rio Grande Valley. As the population increased, the grazing industry emerged and irrigation activities intensified. By 1880, nearly all land

in the Rio Grande Valley that could be irrigated was under development and people there had begun to complain of water shortages (Sorensen and Linford

1 970). By the end of the 19th century, grasslands were severely over grazed, and as a result, stream flows became more erratic, soil erosion was accelerated and the morphologhy of stream channels became wider and shallower. Lee (1907) described the Rio Grande north of El Paso as "mainly a flood-water stream subject to great fluctuations in volume".

Beginning in 1914, vand generally more recently numerous large have ) been constructed in New Mexico on the Rio Grande and its major tributaries

in an effort to alleviate water shortages and to control flooding. These

include, Cabillo, Elephant Butte, Jemez, Cochiti, Gallisteo, McClure, Abiquiu,

El Vado, and Heron. In addition, numerous smaller dams and irrigation diversions have been constructed in the Rio Grande basin in New Mexico.

Water releases from these reservoirs fluctuate severely and downstream flows are often critically depleted.

Membres Drainage - The Mimbres River drainage is an isolated fluvial system of the endorheic Guzman Basin. Perennial flowing water of the Mimbres River exists in two discrete segments. These segments are separated by a 11 km portion of the riverbed that is normally dry.

One segment comprises the headwaters of the drainage, including the forks of the Mimbres River which originate on the western slope of the Black 10" Range of . These forks merge within 5.0 - 6.5 of their origins to-form the main stem of the Mimbres River. The river con- tinues in a southerly direction for approximately 5 km at which point the water becomes subterranean. Within this segment, the Mimbres River is characterized as a high-gradient coldwater stream with moderate flows, alternating between riffle and shallow pool habitats. The downstream 6MA,A4 perennial segment of the Mimbres river extends approximately 40 km and -6.444-s

as surface entity (except during flooding and high runoff) near Dwyer. This

portion of the Mimbres River is characterized as a moderate gradient stream

that flows within a well defined channel with numerous riffles and pools.

Hower

(1857) described the terminus of the Mimbres River in the 1850's as being

a series of pools or lagoons, surrounded by thickets of willows (Salix spp.)

for 4 to 6 miles above this point the river is said to have been up to 2.5

feet deep and to have flowed at 2.5 miles per hour, in the summer. Beginning

in the late 1800's, the river has become wider and shallower--probably

largely as a result of overgrazing (Boles and Dick-Peddie 1983). Today,

the river terminus is usually dry, and ordinarily at high water the stream

averages 13 feet wide and less than a foot in depth. Diversions have

reduced the quantity of water in the river, while uneven flows, flooding,

and repeated stream modifications by local landowners have combined to

eliminate the conditions described by Antisell (op. cit.) to a drastic degree.

Pecos Drainage - The originates in the Sangre de Cristo Mountains

of and extends 920 km SSE to its confluence with the

Rio Grande in west central Texas. Within New Mexico, the Pecos River is

bounded on the east by the , and on the west by a series of mountain ranges, including: the Gallinas, Jicarilla, Capitan, Sacramento and .

The Pecos River upstream from Pecos is a high-gradient, coldwater stream with swift, turbulent flows. Between Pecos and Lake Sumner, the river flows through •a region of rough topography where the river is entrenched and the gradient produces moderate to swift stream flows with numerous riffle areas. Between Lake Sumner and Lake McMillan, the river meanders. through a broad valley of low relief. The river between Lake McMillan and the state line flows through an area of low relief with a "stairstep" stream morphology. Here, stream flows alternate between riffle areas and deep pools. Much of the water in this section, and for some distance downstream

in Texas, is brackish, with salinities as high as 25.1 ppt and frequently averaging greater than 10 ppt (Hubbs 1957, Hubbs and Echelle 1972).

Four major reservoirs presently impound water of the Pecos River in

New Mexico (i.e., Santa Rosa, Sumner, McMillan and Avalon dams). In addition

Red Bluff (in Texas) impounds water of the Pecos River that backs up

into New Mexico, and there are several minor unpoundments in the Carlsbad area. The first of these impoundments (i.e., McMillan Reservoir) was con- structed in 1893. The timing and volume of water releases from these

impoundments are dictated primarily by irrigation demands. During the irrigation season (April-October), releases from these impoundments fluctuate severely and downstream flows are often critically depleted, including by main channel diversions and pumping of ground and surface water. Contrasting this, flows are consistently low during winter months (November-March) when few irrigation releases are made. As a result of this manipulation of the water supply, no flows have been recorded in localized portions of the

Pecos River for 1 02 of each water year (Reiland 1980).

Some insight into the pre-impoundment conditions of the Pecos River is provided by Brevet Captian John Pope (1854) who was exploring a prospective route for the Pacific Railroad near the 32nd parallel. On March 8, 1 854

Captain Pope made the following observation of the Pecos River near its confluence with the :

"The water rushes over "falls", in a bold and rapid

current, at a rate of nine knots per hour. The

river at this point has a good rocky bottom; is

about eighty feet wide, and two and a half feet

deep. There are seven little islands, through which

the stream flows with great force." On March 9, 1 854 Captain Pope made the following observations of the

Pecos River south of the Texas-New Mexico state line:

"The river below the 32nd parallel changes its character

from a rocky bed, with occassional rapids, to soft mud

bottom and banks. 44j4, From the 32nd parallel to its

mouth, the Pecos is always navigable for small stern-

wheeled boats, the difficulties consisting merely in its

extreme crookedness and narrowness. The banks are per-

pendicular, about ten feet high and falling into the

stream constantly - the deep water being uniform from

one shore to the other. The average width would not

exceed 100 feet."

Given this anecdotal evidence it seems apparent that flows Wore volumous prior to the construction of impoundments. Although nothinu can be inferred from these accounts concerning the periodicity and permanicy of these flows, the reported depths would likely have imparted a perennial quality to these habitats even in the most severe drought.

Tularosa Basin - The lies between the Rio Grande and Pecos basins in southcentral New Mexico. It is an with a maximum length of about 150 miles and a maximum width of about 60 miles and an area of approximately 6,000 square miles (Meinger and Hare 1915). The Jicarilla,

Sierra Blanca and Sacramento mountains form the eastern boundary of the basin. Chupadera Plateau along with the Gallinas, Little Burrow, San Andrews and Oregan mountains form the eastern boundary. Mesa Jumanes forms the northern boundary of the basin and a low indefinite divide forms the southern boundary (Meinger and Hare 1915).

Perennial water within the interior of the Tularosa basin is limited to a few isolated springs and seeps along with the outflow from these systems. The water from these springs and seeps is highly alkaline and generally evaporates or becomes subterranean within a few miles of their origins. Several small, perennial streams also exist in a few of

the mountains that border the basin (most commonly along the east side of

the basin).

The arid climate, scaricity of perennial water, and depredation from

Apache Indians prohibited settlement of the Tularosa Basin by Anglo-Europeans until the 1800's, and travel there was generally discouraged for the same reasons. Consequently, an account of the hydrologic conditions at the start of the historic era is not available. However, there is no evidence to suggest that hydrologic conditions at the start of the historic era were markedly different from present conditions. The earliest written account of the hydrology of the basin comes from Gibbs (1870):

"---- there is a valley of some 200 miles long and 20

miles wide, lying between Sierra Blanca and the San Andreas

and Oscuro mountains, in which there is no stream and

only a few alkaline springs and salt lakes or ponds. .

water of a strongly alkali character is obtained by digging

few feet, and around the edges of the district salt marshes

exist . .

Canadian Drainage - The originates on the east slope of the

Sangre de Cristo Mountains in northeastern New Mexico and flows generally southeastward for approximately 205 river miles to its confluence with the Conchas River. From there the Canadian River makes an abrupt turn to the east and continues approximately 107 stream miles before exiting the state midway between the 35th and 36th parallels. Ultimately, the

Canadian River joins the River a of the Mississippi

River.

Most perennial tributaries of the Canadian River in New Mexico rise in mountain ranges along the western edge of the basin. These tributaries include: the Vermejo, Cimarron, and Mora along with Ocate Creek.

Ute Creek is the only principal tributary of the Canadian River that originates from the eastern side of the basin in New Mexico. However,

Ute Creek is ephemeral.

The Canadian River Basin of New Mexico comprises an area of about

1 2,500 square miles and is represented by approximately equal areas of mountainous, plateau and plains regions (Dalrymple 1939). Fluvial systems cus of the mountain$ portion of the basin are characterized primarily as steep gradient cold water streams. The Canadian River and its tributaries are confined to deep canyons within the plateau region, and these canyons become deeper and wider with descent until plains and scattered high mesas become the dominant topography. The river meanders accross the plains in a canyon that is approximately 100 feet deep (Dalrymple 1939). PROCEDURES

Museum records of fish collections from New Mexico offer valuable clues

--of the composition and distribution of the state's native ichthyofauna.

However, most of these collections were made since 1940, after many changes

in the fish fauna had already occurred. To accurately portrait the native

ichthyofauna, museum collection records are presented in the context of

other pertinent information, including: regional distributions of species

of fish, former hydrographic conditions and drainage patterns, fish remains records of fossil fishes from archaeological sites, A , ecological affinities of species, fish

stocking records, patterns of genetic and morphologic variation, select

literature citations, and past climatic conditions.

A map is presented for each species or subspecies of fish that indicates

the locations where specimens of fish have been collected or observed and

where species of fish have been stocked (see Appendix). Nearly 10,000

museum records of fish collections from New Mexico are included in this

analysis. -The maps are provided to aid in the interpretation of these

volumous data. These records date from T851 to the present, and represent

holdings curated at: University, University of New Mexico, o f

—F - University of Kansas, University of Texas at Austin, Baylor University,

Oklahoma State University, Texas A & M University, Arizona State University,

University of Nevada at Las Vegas, University of Michigan, Illinois Natural

History Survey, Tulane University, Academy of Science and the S RYT U. S. Natural History Mus4amm. Distribution records that are based on valid A

& y_j c 5 4.-0_4 v' S 2b voucher specimens are plotted on the maps with small symbols (ex. a) which

contrast the slightly larger symbols (ex. ) that are used to plot distribution records found in the literature and I A

stocking records. This distinction is made to help interpret the origins btA of various species of fish, especially game species. Distrition records based on archaelogical evidence are not plotted on the maps, although they are referred to in the narrative.

Inter- and intrabasin distribution patterns of species of fish are examined in relation to the geomorphic evolution of New Mexico's developing

fluvial systems to determine the probable origins and dispersal routes of

the statds native ichthyofauna. The ecological affinities of species are discussed in relation to historic and prehistoric climatic conditions and patterns of genetic and morphologic variation of select species are discussed where applicable,

The composition of the prehistoric fish fauna is substantiated by paleontological evidence where possible. This paper deals 0.-4( with the post-Miocene ichthyofauna,Ait is fortunate that most of these pre- historic fishes are indistingishable from modern forms and that none are known to be generically distinct from modern forms (C.L. Smith 1 562; Wilson

1 968; Miller 1965). Although few species of fossil fish are known from

New Mexico, several are known from adjacent Texas and along with southwestern Kansas.

The chronology of collection records and other distribution data

is indicated on the maps to reveal changing distribution patterns and as -ic) to provide clues Io4,1the composition of the native fish fauna as it likely appeared at the start of the historic era. Distribution data are arranged within time intervals that represent periods during which sufficient data

exists to form meaningful conclusions about the distribution and status

of each species. Sampling efforts were infrequent prior to 1 95O, however

the fauna is sufficiently represented during the intervals 1850-1900 and 1901-1950. Thereafter, sampling efforts intensif- Lo cif ied and the fauna is -stlf-f-i-e-i-errt-l-t represented during-ni-ne-year-i-n-te-rvola

(i.e., 1951-1960, 1961-1970 and 1971-1980). Finally, diittribution information obtained during 1981 to the present is included in the analysis to present information from several recent intensive surveys for fish. Collections from different time intervals are distinguished by contrasting symbols as indicated by the l egends on each distribution map. Under this system, multiple collections of one species from the same time interval and locality are re- presented on a map by a single data point.

RESULTS AND DISCUSSION

hcAtie —7 Some 51 species of fish are believed tonexist4naturally east of the 4iAx tLx. Continental Divide in New MexicoA(Table 1). These species developed within extensive drainages that were, at various times in geologic history, continuous with major tributaries of the Mississippi and Rio Grande drainages, and to a limited extent, drainage basins of northeretMexico and the .

It appears that most native species of the eastern fish fauna, including their immediate ancestors, gained access to New Mexico's waters no earlier than late Tertiary to mid-Pleistocene time. During this time, the major arrangements of the states complex hydrographic systems were established and uplift of strategic north-south oriented mountain ranges (ex. Sangre de

Cristo, Sacramento and Guadalupe mountains) restricted the east-west dis- persal of fishes, producing faunal patterns reflective of diverse origins.

As a result, species composition differs significantly between the state's eastern drainages and highly endemic faunas have evolved in the Pecos, Mimbres, and Tularosa basins. Table 1. Native fish faunas of major drainages east of the Continental Divide in New Mexico as they likely existed at the start of the historic era (circa 1550 AD). Parenthesis indicate that the status (i.e., native or exotic) is uncertain.

Specific Drainage Rio Grande Taxa Canadian Acipenseridae Scaphirhynchus platorynchus

Lepisosteidae Lepisosteus osseus

Anguillidae Anguilla rostrata

Clupeidae Dorosoma cepedianum

Characidae Astyanax mexicanus X

Cyprinidae Campostoma anomalum - - X X Dionda episcopa X - X - Gila nigrescens X - - - Gila pandora X X (X) Hybognathus nuchalis X X Hybognathus placitus - - - X Hybopsis aestivalis X - X X Hybopsis gracilis - X - X X Notropis amabilis - X - Notropis braytoni X Notropis formosus X Notropis 9irardi - - X Notropis 'emezanus - X X Notropis lutrensis - X X X Notropis orca X - Notropis simus - X X - Notropis stramineus - X X mirabilis - X Phoxinus erythrogaster - - X Pimephales promelas X X X X Rhinichthys cataractae X - X X Semotilus atromaculatus - - - X X

Catostomidae Carpiodes carpio X - X X Catostomus commersoni X X Catostomus plebeius X X Cycleptus elongatus - X X lctiobus bubalus X X Moxostoma congestum X X

Table 1 continued.

Specific Drainage Rio Grande Taxa Canadian Cyprinodontidae Cyprinodon pecosensis X Cyprinodon tularosa X Lucania parva Cyprinodon sp. X

Poeciliidae Gambusia nobilis X Gambusia affinis X X X

Centrarchidae Lepomis cyanellus - X X Lepomis macrochirus - X - X - Lepomis megalotis - - X X

Percidae Etheostoma lepidum X Percina macrolepida X

Salmonidae Salmo clarki virginalis X X (X)

lctaluridae ictalurus furcatus X X ictalurus lupus X lctalurus melas - X lctalurus punctatus - - X Pylodictis olivaris X X

Total number of species per drainage 5 26 1 38 24 4

During middle to late Pliocene time, the Rio Grande flowed south from the San Juan Mountains across the Taos Plateau, then along the east side of the Socorro Valley and through the into south central

New Mexico (Belcher 1975). From there, the Rio Grande flowed toward the

Dona Ana Mountains and through Fillmore Pass into the Hueco and on to the at Presidio (Belcher 1975). During this time, it is likely that Catostomus plebeius or a representative of this phyletic line, entered the ancestrial upper Rio Grande from the southeastern edge of the

Colorado Plateau by way of a middle elevation transfer (in part, Smith 1966).

The ecological affinity of the species for mid-elevation streams prevented expansion of its range over mountains to the east. However, the mountain *Lai- ranges:flank the Rio Grande facilitated its southern range expansion.

During late Pliocene to early Pleistocene the Rio Grande was diverted south from its route through Fillmore Pass and flowed west of El Paso into the playa region of (Strain 1969). It is during this time that

C. plebeius likely gained access to the Guzman basin (including the Mimbres

River).

It is likely that an ancestral form of Salmo clarki virginalis also the gained access to the upper Rio Grande fromAColorado Plateau sometime during middle to late Pliocene time. However, unlike C. plebeius, the ecologic affinity of S. c. virginalis for headwater habitats facilitated its eastward expansion by headwater transfers or captures into the Pecos drainage and possibly the Canadian drainage as well. Subsequent geographic isolation has allowed some genetic differences to accumulate between populations of

S. c. virginalis with broad patterns of genetic variation conforming geo- graphically to the three major drainages of occunence (unpublished data from New Mexico Department of Game and Fish, 1983). It is possible that this species, or its ancestor, entered the Guzman basin in a similar 5

fashion as that proposed for C. plebeius. However, whether or not this occurred is enigmatic. Evidence in support of this stems from a report

by Emory (1848) of a trout from the Mimbres River. •The fact that Emory was able to recognize chubs (Gila) from the Gila River and distingish them

from trout (Salmonidae) adds credibility to this observation. If a native

salmonid was once present in the Guzman basin, it now appears to be extripated

based on the lack of voucher specimens.

During late Pliocene to middle Pleistocene time the ancestral Pecos

River was confluent with the ancestral Rio Conchos, which then drained the

volcanic highlands of northern Mexico and flowed generally east and south- eastward into the (Belcher 1975). During this time the climate of the lower Pecos drainage was much wetter and cooler than present.

The continuity of these systems and the equable climate (compaired to present conditions) enabled several modern species or their progenitors

to ascend the Pecos River into New Mexico, perhaps including: Anguilla

rostrata, Dorosoma cepedianum, Astyanax mexicanus, Dionda episcopa, Ictiobus bubalus, Moxostoma congestum, an ancestral Cyprinodon, and Lucania parva.

The native occurrance of L. parva in the Pecos River of New Mexico is sub-

stantiated by morphometric and meristic evidence (Hubbs and Miller 1965);

the native occurrance of the other species is substantiated by early collection

records (see appendix) and conformation with regional distribution patterns.

In addition, Dorosoma petenense gained access to the lower Pecos drainage

during this time as is evident from a fossil of this species found in late

Pliocene deposits near Loving, New Mexico (Miller 1982). However, the arid conditions of Holocene time, coupled with cold winters likely eliminated

this species in New Mexico prior to historic times.

The progenitor of Gambusia nobilis likely gained access to the ancestral

Pecos River during late Pliocene to middle Pleistocene time. Gambusia

nobilis evolved from a highly variable and widespread progenitor which also 6 gave rise to five other modern species of Gambusia that presently occur allopatrically, for the most part ) in various waters from south central

Texas to northeastern Mexico (Hubbs and Springer 1 957). Gambusia nobilis, as well as other contemporary species of this progenitor, evolved in

isolated spring habitats. Populations that once occurred in intervening waters were apparently victimized by drought conditions of an interglacial __ — period along with subsequent competition and hybridization with a ffinis, This latter species entered the wge of G. no ilis A during an i'rrteryldLldl period from the east

(Hubbs and Springer 1 957).

During middle Pleistocene time the Rio Grande was diverted east from

its course into northern Mexico to a route through El Paso Canyon into the

Hueco Bolson (Belcher 1 975). From there the Rio Grande again gradually worked southward toward Prisidio, Texas where it joined the ancestral Rio

Conchos (Belcher 1975). .140-01-er diversion of the Rio Grande from northern

e Mexico 40§....G43ffir+etik, the ingress and egress of fishes to the Guzman basin

404-affml-4444-ric44141, resulting in a highly endemic and unsaturated fauna. ilimw-i-erg-this time, Notropis formosus (c,14-al-y-44-64-ed Notropis lutrensis A Contreras-Balderas 1975, 1978) and the progenitor of Gila nigrescens, likely gained access to the Guzman basin from the Rio Yaqui drainage. This is believed to have occured by way of a low elevation headwater transfer, perhaps via the headwaters of the Rio Papigochic, near Migaca, , which may have been confluent at one time with the Rio of the Guzman drainage (Miller 1 958). It is likely that an ancestral Cyprinodon and P. promelas, of southern and eastern affinity respectively, gained access to the Guzman basin in the same manner.

It has been suggested that G. nigrescens and Gila pandora were derived

from a common ancestor from the Yaqui basin (Rinne 1969, Uyeno 1961). Diff- - — erentiation between these species occurred as a result of geographic isolation, 7 with G. nigrescens becoming restricted to the Guzman basin while G. pandora gained access to the Pecos drainage and ancestral lower Rio Grande via the

Rio Papigochic when it was confluent with the Rio Conchos (presumably some- time during middle Pleistocene time or more recently).

Once the Rio Grande joined the ancestral Rio Conchos at Presidio,

Texas, (during middle Pleistocene time) integration into the present day system was complete. This enabled several faunal elements, to ascend the

Rio Grande into New Mexico, perhaps including: Scaphirhynchus platorlynchus,

A. rostrata, D. cepedianum, A. mexicanus, D. episcopa, G. pandora, I. bubalus,

M. congestum, and G. affinis. The native occurrance of these species in the Rio Grande of New Mexico is substantiated by early collection records

(see appendix) and conformation with regional distribution patterns.

Major mountain ranges that presently lie between the Pecos River and the Rio Grande (ex. Sangre de Cristo and ) were uplifted during late Pliocene time (Belcher 1975). These events deprived the ancestral

Pecos River of much of its headwaters because they were diverted into the newly established south-flowing Rio Grande. It is deduced that the Pecos

River of late Pliocene time headed within the approximate area that presently comprises the Tularose basin. From there the river flowed along the east side of the Sacramento upwarp, and across the Stockton- to join the ancestral Rio Grande near the present mouth of the Pecos River

(Belcher 1971). This drainage provided a likely immigration route to the

Tularosa Basin for an ancestral Cyprinodon that appears to have been wide- spread at this time throughout the lower Pecos drainage. Uplift of the

Guadalupe and Sacramento mountains during late Tertiary to middle Pleistocene time "beheaded" the ancestral Pecos River in the area of Sierra Blanca and may have diverted the headwaters of this system into the endorheic Tularosa basin. Eventually, differentiation of the ancestral Cyprinodon occurred as 8a a result of geographic isolation, producing the contemporary form: Cyprinodon tularosa.

Conclusive evidence is laking of other species of fish gaining access to the Tularosa basin. Behenke (1981) cites evidence of a possible headwater transfer of S. c. virginalis from the or (of the

Pecos River basin) to Indian Creek (of the Tularosa basin). However, whether or not this occurred is unknown. I f other species of fish did exist in the Tularosa basin, it is likely that they were eliminated there as a result of drought conditions brought on by tetonic activities that deprived the basin of water and increasingly arid climatic conditions of Holocene time.

Following the uplift of the Guadalupe and Sacramento mountains, the beheaded ancestral Pecos River became a headward eroding stream which slowly worked its way north and eastward (Thomas 1972). In the process, the Pecos

River captured several streams during late Pleistocen time that once drained eastern New Mexico, including the of , and the upper ancestral whcih occupied a valley in the present position of the upper Pecos River above Fort Sumner (Baker 1916, Thomas 1972). It is likely that Etheostoma lepidum and Percina macrolepida gained access to the

Pecos River through the Colorado River when these systems were confluent.

Capture of the ancestral Brazos River by the Pecos River allowed the infusion of several Mississippian species into the Pecos River, including: Lepisosteus osseus, Hybognathus nuchalis, Notropisjeutrensis, Pimephales promelas, 7 Carpiodes carpio, Cycleptus elongatus, Lepomis macrochirus, Lepomis megalotis,

Ictalurus furcatus and Pylodictis olivaris. The progenitors of several other modern species likely gained access to the Pecos River by this same means, including that of: Notropis amabilis, Notropis braytoni, Notropis

emezanus (all sibling species of Notropis atherinoides); the Notropis simus- orca complex (closely related to species of Alburnops); and I ctalurus lupus

(sibling species of Ictalurus punctatus). The native occurrance of these 8h species in the Pecos River of New Mexico is substantiated by early collection records (see appendix) and conformation with regional distribution patterns.

Echelle and Echelle (1978) suggested that the progenitor of Cyprinodon pecosensis was introduced into the Pecos River from the Brazos River when these drainages were confluent (e.e., during late Pleistocene time). However, the Brazos River seems an unlikely source for this ancestral Cyprinodon that exists in view of the hiatus in the distribution of congenersAthere

and in parallel drainages such as the Red and Colorado . Only

two species of Cyprinodon occur naturally in these drainages: Cyprinodon

rubrofluviatilis exists naturally only in the headwaters of the Brazos and Red rivers (Echelle and Echelle 1978), whereas Cyprinodon variegatus

exists naturally only in coastal waters (ex. of the Gulf of Mexico) and

in esturine habitats.

The more logical origin of the progenitor of C. pecosensis is from

the south. As previously discussed, an ancestral Cyprinodon that gave

rise to C. tularosa is believed to have been present in the lower Pecos basin during late Pliocene time. This same ancestor likely gave rise to

three other species of Cyprinodon in the Pecos basin that are extant:

C. ecos s, C. bovinus and C. elegans. Cyprinodon pecosensis is wide-

spread in the Pecos drainage ■)1: Jew Mexico and Texas, whereas C. bovinus and C. elegans are represented by relictual, allopatric populations in

Texas. Differentiation of these three forms occurred as a result of geographic isolation; however, limited contaot and genetic introgression between C. pecosensis and C. bovinis is suspected (Echelle and Echelle 1978).

By late Pleistocene time, the geomorphic evolution of the Pecos River,

together with favorable hydrologic and climatic conditions, enabled the progenitor of C, pecosensis to ascenci the river to the vicinity of Fort Sumner, New Mexico. From there, the species gained access to the

Brazos River when the headwaters of that system were captured by the Pecos

River (see previous discussion). Differentiation of this ancestral S kbsp f„s y

Cyprinodon ocurred as a result of geographic isolation in the Brazos drainage i AI Re. • k producing the contemporary form:C. rubrofluviatilis.

During late Tertiary to middle Pleistocene time, the upper Canadian River 1 0

drained into the Pecos River in the vicinity of Ft. Sumner. Presumably

during late Pleistocene time (or maybe more recently), a headward-eroding

lower Canadian River captured the ancestral "Canadian" tributary of the

Pecos River, thereby establishing the Pecos and Canadian rivers in their

present courses (in part, Thomas 1972). Once continuity of the upper

Canadian River with extensive eastern drainages had been established

numerous species of fish were able to ascend the river into New Mexico,

including: A. rostrata, D. cepedianum, Campostoma anomalum, Hybognathus placitus, H. aestivalis, Hybopsis gracilis, Notropis girardi, N. l utrensis,

Notropis stramineus, Phenacobius mirabilis, Phoxinus erythrogaster, P. promelas, Rhinichthys cataractae, Semotilus atromaculatus, C. carpio,

Catostomus commersoni, Fundulus zebrinus, G. affinis, Lepomis cyanellus,

L. megalotis, lctalurus melas, and lctalurus punctatus.

The native occurrence of these species in the Canadian drainage of

New Mexico is substantiated by early collection records. In addition eight of these are known from fossils recovered from Illinoian deposits in north- western Oklahoma and southwestern Kansas (Cross ). The fossil record from these deposits also suggest that several other species of fish may have gained access to the Canadian drainage, including: L. osseus, Notemigonus crysolucas, Lepomis humilis, Micropterus salmoides and Perca flavescens

(Cross ). However, the chronology of collections do not substantiate the natural occurrance of these species in the New Mexico portion of the

Canadian River during late Tertiary to middle Pleistocene time, they likely became extirpated there as a result of reoccuring drought conditions of Holocene time.

It is possible that Morone chrysops also gained access to the Canadian

River into New Mexico during late Tertiary to middle Pleistocene time. The species is native to the lower (Hubbs 1982) and presumably it was native in adjacent drainages (ex. the Canadian drainage). The wetter and cooler climatic conditions of this time may have enabled the species 11

to ascend the Canadian River. However, the chronology of collections do not

substantiate the natural occurrance of this species in the Canadian drainage

of New Mexico during historic times. If the species did ascend the Canadian

River during late Tertiary to middle Pleistocene time, it likely became

extirpated there as a result of arid conditions of Holocene time.

During late Pleistocene or early Holocene time, the Conchas River

(a tributary of the Canadian River) appears to have captured a tributary

of the Gallinas River (of the Pecos basin) thereby allowing transfer of

faunal elements between these drainages. Among the species of fish that

may have gained access to the Pecos drainage during this time are several

that were, up to that time, unique to that drainage, including: C.

anomalum, H. aestivalis, H. gracilis, N. stramineus, Phenacobius mirabilis,

R. cataractae, S. atromaculatus, C. commersoni, F. zebrinus, and L. cyanellus.

At the same time, G. pandora may have gained access to the Canadian drainage.

Culaea inconstans is known from the Vermejo River drainage in north-

eastern New Mexico, with populations occurring in Merrick Lake, the Vermejo

River and Stubblefield Lake. This species was first collected in New Mexico

by W. Wolfrum in Merrick Lake on 17 April 1954. The New Mexico populations

are separated geographically by approximately 900 air kilometers from the

next closest population, which is in the vicinity of Omaha, Nebraska. As

such, New Mexico populations of the species are of special zoogeographic

interest. It is possible that C. inconstans invaded the Canadian basin and ultimately the Vermejo drainage during late Pleistocene time when life

zones were shifted 1,200 m lower than their present position (Martin 1963) and alternating glacial and pluvial conditions prevailed. Theoritically,

under these conditions, C. inconstans could have occupied lower elevations

and more southern latitudes, permitting access to the Vermejo drainage

from the lower Mississippi drainage. The ensuing warmer and dryer climates of Holocene time made the waters of southern latitudes unsuitable 1 2

for the species except for the Vermejo drainage in New Mexico where a

relictual population remained. In spite of this speculation, the New

MexicOopulations are believed to be exotic occurances because they do

not conform to the pattern of morphometric and meristic clinal variation

observed elsewhere for the species, as reported by Nelson (1969). The

late discovery of C. inconstans in New Mexico and the popularity of waters

in the Vermejo drainage with anglers suggests that the occurances of this

species in the Vermejo drainage are the result of a bait bucket introduction

or an introduction incidental to stocking an exotic salmonid. The New Mexico

stock is suspected to have originated in or Wisconsin.

As previously discussed, 26 species of fish of eastern origin gained

access to the Pecos River of New Mexico. Of these species, only 15 appear

to have gained access to the Rio Grande of New Mexico, including: L. osseus,

H. nuchalis, H. aestivalis, H. gracilis, N. jemezanus, N. l utrensis, the

ancestral form of the N. simus-orca complex, P. promelas, R. cataractae,

C. carpio, C. elongatus, L. macrochirus, I. furcatus, and P. olivaris.

Cycleptus elongatus and L. osseus are known from the Rio Grande of New Mexico

only by their skeletal remains, which were recovered from Pueblo Indians

ruins at Bandelier National Monument and Chaco Canyon, respectively (Gehlbach and Miller 1961); the native occurrance of the other species in the Rio Grande of New Mexico is substantiated by early collection records (see appendix).

By necessity, these species gained access from the Pecos River to the

Rio Grande of New Mexico by way of the present confluence of these drainages.

It is inferred from the ecological requirements of most of these species that diipersal to the Rio Grande occurred primarily during the Pleistocene era when life zones were shifted 1,200 m lower than their present position

(Antevs 1928) and pluvial conditions prevailed. It is likely that these transfers occurred approximately 20,000 years ago when the last major pluvial maximum occurred in the south west (in part, Martin 1963; Geol.). 1 3

However, it is possible that any number of these species were transferred more recently because pluvial conditions persisted until 12,000 B. P.

(in part, Martin 1963; Geol.). LITERATURE CITED

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Belcher, R. C. 1975. The geomorphic evolution of the Rio Grande. Baylor Geological Studies Bull., 29: 1-64.

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