Horned Lizards by Eric R. Pianka and Wendy L. Hodges Erroneously called "horny toads," horned lizards are bizarre, spiny, ant-eating lizards unlike any other lizards in North America. Fourteen species are currently recognized, 8 of which are found within the continental USA (one reaches southern Canada), and 6 other little-known species are restricted to Mexico (one reaches Guatemala). Most horned lizard species are well represented in the fossil record by the Pleistocene (1 million years ago, mya), P. cornutum is found in the upper Pliocene (3 mya), and P. douglasi is known from the mid Miocene (15 mya). Three species went extinct in the late Pliocene-early Pleistocene, long before there were any humans. The genus is thought to have split from an ancestor shared with the sand lizards (Uma, Callisaurus, Cophosaurus, and Holbrookia) during the late Oligocene-early Miocene (23- 30 mya). Humans and horned lizards have shared each other's company for thousands of years. This relationship is recorded from Anasazi, Hohokam, Mogollon, and Mimbres cultures through their use of horned lizard images on pottery, petroglyphs, effigy bowls, figures, and shells. Hopi, Navajo, Papago, Pima, Tarahumara and Zuni cultures portray horned lizards in their ceremonies and stories as symbols of strength. Piman people believe horned lizards can cure them of a staying sickness by appealing to the lizard's strength and showing their respect to the animal. They formulate a cure by singing at a patient's side songs describing the lizards and their behaviors. A horned lizard fetish may be placed on an afflicted person's body during the songs. Native Mexican people also respect horned lizards attributing the words, "Don't tread on me! I am the color of the earth and I hold the world; therefore walk carefully, that you do not tread on me." A Mexican common name for horned lizards is "torito de la Virgen" or the Virgin's little bull. This name apparently was given to the lizards both because of their horns and because horned lizards are sacred to many people due to their blood squirting behaviors, otherwise considered weeping tears of blood. These interesting lizards were first introduced to European audiences in 1651 by the Spaniard Francisco Hernandez. Hernandez was fortunate to observe a living individual which squirted blood from its eyes -- he noted this behavior in his report on the first scientific expedition to Mexico by Spain. Over a century later in 1767, a Mexican cleric of Spanish descent, Clavigero, also showed his wonder of horned lizards in his illustrated volumes of Mexican history. Still later, in 1828 Wiegmann coined the official scientific generic name Phrynosoma, which is Greek for toad-bodied (phrynos means "toad", soma means "body"). Species descriptions Horned lizard species are distinct and easily recognized from each other. Arrangement of occipital and temporal horns on the head are enough to distinguish species, but other features such as number of rows of lateral, abdominal fringe scales and dorsal scale patterns are helpful, too. Color or color patterns generally are not good distinguishing features because these lizards are extremely variable and tend to match the color of the sand or rocks in their local environment. However, color can be used in certain circumstances as stated in species' descriptions below. The geographic ranges described below are based on historic records and do not reflect species' current distributions. Ranges of most species have been severely reduced. Phrynosoma asio, Long Spined Horned Lizard Diagnostic Characters: Two rows of abdominal fringe scales Three rows of large conical scales widely separated on the dorsum Gular scales arranged in serrated longitudinal series Large keeled ventral scales numbering 30-35 across the widest point constitute a formidable armor. Geographic Distribution: This is the largest species attaining snout-to-vent length (SVL) of 115 mm and total length of 202 mm (8 inches). It occurs in southern Mexico from Colima through coastal Michoacan, Guerrero, Oaxaca, to Chiapas, and in the Balsas Basin. It is recorded from Guatemala. Phrynosoma boucardi Diagnostic Characters: Two long occipital spines that project vertically Temporal spines relatively long, projecting horizontally Superciliary region elevated Weakly keeled gular scales While this species has been recognized as distinct, it has also been relegated to the subspecies P. orbiculare boucardi because the morphological characters that diagnose it are found on intermediate forms in the P. orbiculare group, although not all characters are found together. P. orbiculare is a wide ranging species and occurs very near specimens called P. boucardi More information is needed to sort out the taxonomy of P. boucardi. Phrynosoma braconnieri Diagnostic Characters: One row of lateral abdominal fringe scales Two moderately developed occipital spines, short, heavy, separated Temporal horns very reduced Very short tail All ventral scales keeled Nostrils in the line of the canthus rostralis Geographic Distribution: This very rare horned lizard occurs in pine-oak woodland and xeric thorn-scrub of Puebla and Oaxaca, Mexico. It may inhabit Veracruz. Phrynosoma cornutum: Texas horned lizard Diagnostic Characters: Single pair of occipital spines Two rows of lateral abdominal fringe scales Enlarged modified dorsal scales with 4 distinct keels Single row of enlarged gular scales Keeled non-mucronate ventral scales Postrictal scale absent White middorsal stripe Geographic Range: From central Kansas, extreme southwestern Missouri, and the southeastern corner of Colorado southward and westward throughout most of Oklahoma and Texas (including coastal barrier islands), southeastern half of New Mexico and southeastern corner of Arizona to Mexican states of Sonora, Chihuahua, Durango, Coahuila, Nuevo Leon, Tamaulipas, San Luis Potosi, and Zacatecas. Phrynosoma coronatum: Coast horned lizard Diagnostic Characters: Two rows of lateral abdominal fringe scales Two occipital spines, 3-4 times longer than basal width, not in contact 4-5 temporal spines Smooth ventral scales Three or more rows of enlarged gular scales Mate belly to belly Geographic Range: From central California, west of the Sierra Nevada south throughout southern California, west of the Mojave desert, throughout Baja California, west of the Sierra Juarez and Sierra San Pedro Martir, Mexico. Up to six subspecies of P. coronatum have been recognized throughout its range. One subspecies has also been named a unique "species" (its taxonomic status is in doubt), P. cerroense, which occurs on the Pacific island, Isla de Cedros. Phrynosoma ditmarsi: Rock horned lizard Diagnostic Features: Occipital and temporal horns reduced to flaring expansions Deep and narrow occipital notch High post-orbital ridge Large vertical expansion of the mandibles Bare tympanum in the anterior neck fold posterior to a vertical row of four spines One row of lateral abdominal fringe scales surrounded by prominent scales Geographic Range: Northern Mexico at rocky sites in oak and oak-pine woodlands and short-tree Sinaloan deciduous forest along the western aspect of the Sierra Madre Occidental in the State of Sonora, north of Yecora, at elevations of 1050-1425 meters. This species was "lost" to science for about 65 years. Its unique habitat preferences and limited distribution, combined with a very imprecise holotype locality record made it difficult to locate. An extraordinary effort by Vincent Roth based on a cross-correlational analysis of gut contents from only three specimens led to its rediscovery. Phrynosoma douglasi: Pygmy Short-Horned Lizard Diagnostic Characters: Specimens are very small in size One row of lateral abdominal fringe scales Head spines very short and reduced to tubercles, project vertically Deep notch between occipital spines Dorsal scales irregular in size and distribution, set in a rosette of smaller keeled scales Geographic Range: This species is restricted to the Pacific Northwest in northern California, north through Washington to British Columbia, Canada and east through southeastern Idaho. Restricted to higher elevations. Phrynosoma hernandezi: Short-horned lizard Diagnostic Characters: One row of lateral abdominal fringe scales Reduced occipital spines directed posteriorly, parallel with temporal spines Dorsal scales arranged in 6-8 longitudinal rows, rosettes absent or poorly developed. Geographic Range: Wide ranging, P. hernandezi is found from southern Alberta, Canada, through Montana, Utah, Colorado, Arizona, New Mexico and into Mexico through northeastern Sonora, Chihuahua, and Durango. Restricted to higher elevations. Until recently, this species was previously included with P. douglasi. Phrynosoma mcallii: Flat-tailed horned lizard Diagnostic Characters: Flat and broad tail Two or three rows of lateral abdominal fringe scales Two elongate, sharp occipital horns 3-4 times longer than the basal horn width Six temporal horns Nostrils inside canthus rostralis Tympana are not externally visible Enlarged lateral row of gular scales Olive or brown mid-dorsal stripe Geographic Range: The original historical range is recognized as extending from the Coachella, Imperial, and Borrego Valleys in Riverside, Imperial, and extreme eastern San Diego Counties, California; west of the Gila and Tinajas Altas Mountains and south of the Gila River, Yuma County, Arizona; northeastern Baja California, east of Sierra de Juarez and north and west of Bahia de San Jorge in Sonora, Mexico. The distribution of this species is now much more restricted. Phrynosoma modestum: Round-tailed horned lizard Diagnostic Characters: Tail round in cross section No lateral abdominal fringe scales Four occipital horns short and equal in length Ear concealed in some, exposed in others Chin shields contact infralabials throughout Solid, pale coloration, white, tan, or pink Dorsal marks restricted to dark neck patches, dark tail bands, some shadowing around limbs Unique rock mimicry behavior Geographic range: Western half of Texas, north to Albuquerque, New Mexico, west to southeastern Arizona, south through northeastern Sonora, most of Chihuahua, through the northern third of Durango, to northern Zacatecas and northeastern Nuevo Leon, Mexico. Phrynosoma orbiculare: Chihuahua desert horned lizard Diagnostic Characters: One row of lateral abdominal fringe scales Two short occipital horns Temporal horns extend nearly as far as occipital horns Smooth ventral scales Very short tail Geographic Range: This wide ranging Mexican species is found on the northern plateau from Chihuahua, Durango, Nuevo Leon, southeast through Morelos, Puebla and Veracruz. Desert horned lizard, Phrynosoma platyrhinos. Phrynosoma platyrhinos: Desert horned lizard Diagnostic Characters: One row of lateral abdominal fringe scales Two moderately elongated occipital horns, not in contact at base Enlarged chin shields Nostrils inside the canthus rostralis Blunt snout Geographic Range: Lowland deserts in southeastern Oregon, southwestern Idaho through Nevada, western Utah, California, Arizona, south barely into northern Baja California and northern Sonora, Mexico. Phrynosoma solare: Regal horned lizard Diagnostic Characters: One row of lateral abdominal fringe scales Four large occipital horns, in contact at base and continuous with six temporal horns forming a large crown of ten horns. Nostrils inside the canthus rostralis Ventral scales keeled Geographic Range: The species is restricted to Sonoran desert in south-central Arizona, east to extreme southwestern New Mexico, south through most of Sonora (including Isla Tiburon) and into northern Sinaloa, Mexico. Phrynosoma taurus: Diagnostic Characters: One row of lateral abdominal fringe scales Occipital spines reduced and separated by a notch Temporal area greatly enlarged posterolaterally, ending with two heavy, moderately long spines All ventral scales keeled Nostrils inside the canthus rostralis Geographic Range: Little is known about this species. It is found in montane chaparral-oak forest and desert areas of the Sierra Madre del Sur, south and southeast of Mexico City, in Guerrero and Puebla, Mexico. Geographic distributions of egg laying species. Geographic distributions of live-bearing species. Phylogeny and Natural History Phylogeny allows us to trace evolutionary history and relationships of organisms. Much like humans draw their genealogies, or family trees, to discover where their blue eyes or baldness came from, or perhaps whether they are genetically predisposed for cancer -- systematists construct such trees to show how different species have evolved. Ecologists use phylogenetic relationships to learn how characteristics of species evolved, or how different species acquired traits and evolved to occupy their current niche. Principles of parsimony are used to identify the simplest explanations for how a trait evolved. There are two major lineages of horned lizards, one of which lay eggs (oviparous) while members of the other group give birth to living young (viviparous). Although the ancestral state is oviparity, one lineage of horned lizards, all high altitude species, has evolved live bearing (braconnieri, boucardi, ditmarsi, douglasi, hernandezi, orbiculare, and probably taurus). Viviparity appears to have arisen only once in the genus, rather than independently 5 times. Interestingly, all species are montane which provides support for the idea that drier and colder mountain climates demand that montane lizards retain their progeny internally until birth rather than laying eggs. Two hypothetical phylogenies for horned lizards. Horned lizards are a rather fecund group, and lay or give birth to many offspring compared to other lizards. The median clutch size for P. cornutum is 25 (one specimen laid 40 eggs!), P. asio lays 17 on average, and P. hernandezi bears up to 16 live lizards. Reproductive effort measures the resources given to producing offspring and is often measured by comparing the weight or volume of the offspring to female volume or body weight (relative clutch mass, or RCM). RCM among horned lizards ranges from 13% to 35%, (offspring constitute from 13-35% of a female's weight). Females can have a few large versus many small progeny. Some species also reproduce twice in a season. This large investment in offspring throughout the active season weighs down females and makes them vulnerable to predators. Because babies are tiny and easy prey for a multitude of predators, horned lizards would go extinct without such high fecundities. An interesting more recent analysis of horn lengths among horned lizards is shown in the above figure from Leache and Maguire (2006). Preferred phylogeny for Phrynosoma based on combined mtDNA and nuclear data. The most parsimonious reconstruction of antipredator blood-squirting (ABS) is mapped on the phylogeny (black bars). Blood squirting behavior was ancestral and has been lost 4 times. Silhouettes of Phrynosoma heads are shown to illustrate the variation in cranial horn anatomy, color coded to correspond to relative horn length (ancestral state reconstructions mapped on each node). The asterisk adjacent to P. taurus indicates that the effective length of the squamosal horns of this unusual species are longer than portrayed by the measuring technique employed. Four Phrynosoma clades, TAPAJA, ANOTA, DOLIOSAURUS, and BREVICAUDA are marked (from Leache and Maguire 2006). Horned lizards have evolved a variety of mechanisms to avoid their predators which include loggerhead shrikes, hawks, roadrunners, a variety of snakes, coyotes and foxes. Their first line of defense is to remain cryptically hidden from a predator's sight. This is accomplished by three things, matching the background color of the substrate, possessing various spines and fringes of scales which decrease their shadows, and they remain motionless when approached. Secondly, their formidable body armor of spines and horns pose a significant threat to many predators as witnessed by snakes and birds found dead with lizards' horns projecting through predators' throats. Horned lizards will capitalize on their armor by inflating their bodies with air until they look like spiny balloons. At least four species of horned lizards (but not all species), coronatum, cornutum, orbiculare and solare, squirt blood from their eyes when attacked, especially by canine predators such as foxes and coyotes. The canine will drop a horned lizard after being squirted and attempt to wipe or shake the blood out of its mouth, clearly suggesting the fluid has a foul taste. Older horned lizard phylogenies suggest that blood squirting behavior either evolved independently 4 times or it evolved only once and was then lost in subsequent lineages. Leache and Maguire's more recent tree suggests blood squirting behavior was ancestral and has been lost 4 times. This behavior is currently being investigated by Dr. Wade C. Sherbrooke, Director of the Portal Research Station of the American Museum of Natural History. Horned Lizards, continued.