Diversity
The genus
Lampropeltis
exhibits a remarkable diversity across its range, which spans the United States,
Mexico, and parts of Central and South America. This diversity is not only geographical
but also manifests in the wide array of colorations and patterns displayed by different
species within the genus. From the banded or striped black and white of the California
Kingsnake to the red, black, and yellow bands mimicking venomous counterparts seen
in the Scarlet Kingsnake, each species showcases unique adaptations to their specific
habitats and ecological niches. Such variation extends beyond mere appearance; it
reflects the evolutionary history of
Lampropeltis
as these snakes have adapted to a variety of environments, from forests and grasslands
to desert regions. This adaptability and the resulting diversity make the genus
Lampropeltis
a fascinating subject of study for herpetologists and a favorite among reptile enthusiasts.
The mimicry of venomous snakes by certain
Lampropeltis
species, for example, is a sophisticated survival strategy that highlights the complex
interactions between species and their environments. Thus, the diversity within
Lampropeltis
is a vivid testament to the power of natural selection and evolutionary adaptation.
Geographic Range
Lampropeltis
species span a significant portion of the Americas, from the Nearctic region down
through the Neotropical region, showcasing their adaptability to a wide range of environmental
conditions. Their presence in these regions underscores their ecological versatility,
from temperate zones in the north to the tropical climates in the south.
- Biogeographic Regions
- nearctic
- neotropical
Habitat
Lampropeltis
species exhibit an extraordinary adaptability to a wide range of terrestrial habitats
across the Americas. These non-venomous snakes thrive in environments ranging from
dense forests and woodlands, where fallen logs and dense underbrush provide shelter
and abundant prey, to the open landscapes of grasslands and prairies, where they utilize
burrows and rock piles for protection while hunting small mammals. In arid deserts,
they cleverly use rock crevices and underground burrows to escape extreme heat, adapting
their foraging habits to the cooler times of day to hunt desert-dwelling creatures.
Wetland areas offer them aquatic vegetation and fallen trees as concealment, with
a diet enriched by fish, frogs, and aquatic rodents. Remarkably, they also navigate
human-modified landscapes such as agricultural and suburban areas, exploiting buildings
and gardens for shelter and feeding on the high rodent populations these areas tend
to support. In rocky outcrops and mountainous regions, they find refuge in crevices
and adapt their diet to the local fauna, demonstrating a broad elevational range from
sea level to several thousand feet. This wide habitat utilization showcases their
ecological versatility, relying on specific habitat features like appropriate sheltering
spots, diverse foraging opportunities, and optimal conditions for thermoregulation
to survive and thrive across their expansive range.
- Habitat Regions
- temperate
- tropical
- terrestrial
- Terrestrial Biomes
- desert or dune
- savanna or grassland
- chaparral
- forest
- rainforest
- scrub forest
- mountains
Systematic and Taxonomic History
The evolutionary understanding and classification of the genus
Lampropeltis
, which includes the widely recognized kingsnakes and milk snakes, have undergone
significant transformations over time, especially with the advent of genetic and molecular
research methodologies. Initially, the evolutionary relationships and taxonomy within
Lampropeltis
and between it and other taxa were predominantly inferred from morphological characteristics,
leading to broader, less precise groupings based on physical traits such as scale
arrangements and color patterns. However, recent genetic studies have provided a more
refined perspective, revealing a clearer delineation of
Lampropeltis
unique lineage within the
Colubridae
family and its closer evolutionary ties to other non-venomous snake genera such as
Pituophis
,
Pantherophis
and
Pituophis
. These genetic insights have prompted a reevaluation of species classifications within
the genus, distinguishing more distinct species and subspecies based on DNA evidence
rather than morphology alone. This shift has not only clarified the evolutionary pathways
and relationships within
Lampropeltis
but also led to reclassifications and, in some cases, the merging or splitting of
species or subspecies. While the genus name
Lampropeltis
has remained stable, the deeper, genetically informed understanding of these snakes'
taxonomy reflects a broader trend in herpetology toward more nuanced and accurate
representations of evolutionary relationships, even though individual species and
subspecies within the genus may have experienced name changes or reclassifications
as a result of these advances.
Physical Description
Adults of
Lampropeltis
species typically range from 3 to 6 feet in length and possess a slender, muscular
body with a slightly distinct head and round pupils, indicative of their non-venomous
nature. Their smooth, shiny scales come in a spectrum of colors including black, white,
yellow, red, and brown, adorned with patterns that can be bands, stripes, blotches,
or combinations thereof. The diverse colorations and patterns, which vary significantly
across species, serve not only as camouflage but also as a mimicry defense mechanism
against predators. The specific physical characteristics can vary widely from one
species of
Lampropeltis
to another, but these general traits are common among them.
- Other Physical Features
- ectothermic
- heterothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes alike
Development
Lampropeltis species generally have straightforward developmental patterns typical of many reptiles. Eggs are laid by the female and undergo direct development; there is no larval stage as seen in some other reptiles like amphibians. The juveniles hatch fully formed, resembling smaller versions of the adults, and grow larger through a series of ecdyses (shedding of the skin). Lampropeltis species typically exhibit genotypic sex determination (GSD), where sex is determined at fertilization by genetic factors.
- Development - Life Cycle
- indeterminate growth
Reproduction
Lampropeltis
mating behaviors are marked by solitary and opportunistic interactions rather than
complex social structures or long-term pair bonds. The primary mode of finding and
attracting mates relies on chemical communication, with females emitting pheromones
to signal readiness for mating, and males utilizing their keen sense of smell to track
these cues. Visual signals play a lesser role in these initial interactions. During
the breeding season, males may engage in non-lethal combat, a form of wrestling designed
to demonstrate dominance and fend off competition for mating rights. This behavior,
while indicative of polygynous or polyandrous systems where individuals may mate with
multiple partners, does not contribute to a defined social hierarchy or structure
outside of this context. Mating involves the alignment of male and female bodies,
with copulation potentially lasting several hours... The timing of the breeding season,
typically in spring following hibernation, ensures that offspring are born at a time
that maximizes their survival prospects, allowing them to grow before facing the challenges
of their first winter. Thus, the mating behaviors of
Lampropeltis
, characterized by solitary interactions, chemical communication, and the occasional
display of physical dominance, reflect an adaptation to their environments and life
cycles, emphasizing survival and reproductive success without the formation of enduring
social bonds or structures. Since they ley eggs, they are oviparous animals. They
mate once a year and they lay a clutch size of 2-17 eggs.
- Mating System
- polygynandrous (promiscuous)
The reproductive cycle of
Lampropeltis
species, including kingsnakes and milk snakes, is intricately tied to seasonal patterns,
with breeding occurring in spring following hibernation to take advantage of favorable
environmental conditions. Females lay between 2 to 24 eggs, depending on the species
and individual condition, within weeks after mating. There's no traditional gestation
as seen in mammals, but rather an incubation period for the eggs ranging from 40 to
60 days in protected environments that ensure optimal temperature and humidity. Upon
hatching, the young snakes are immediately independent, equipped with a temporary
yolk sac for nutrition and capable of hunting small prey.
Lampropeltis
reach sexual maturity at around two to four years of age, a timeline that varies
by species size and environmental factors. This reproductive strategy, marked by a
lack of parental care post-egg-laying and immediate offspring independence, underscores
the adaptability and resilience of
Lampropeltis
species in their natural habitats, ensuring the survival and dispersal of the species
across their wide geographic range.
- Key Reproductive Features
- iteroparous
- year-round breeding
- sexual
- fertilization
- oviparous
In
Lampropeltis
species, parental investment is minimal and primarily focused on the pre-hatching
stage, with the female's role being crucial in selecting an appropriate nesting site
for her eggs. This choice represents the extent of parental care, aiming to optimize
the microenvironment for egg incubation, thereby indirectly offering protection from
predators and environmental hazards. Post-fertilization, there is no further investment
from either parent, as males and females part ways immediately after mating. Upon
hatching,
Lampropeltis
offspring are entirely independent, equipped with a yolk sac for initial nourishment
but otherwise capable of fending for themselves without any parental guidance or support.
This reproductive strategy highlights the solitary nature of
Lampropeltis
species, where the survival of the young is ensured through the mother's selection
of a secure and conducive nesting location, rather than through direct parental care
or provisioning.
- Parental Investment
- no parental involvement
- precocial
Lifespan/Longevity
Lampropeltis
species exhibit notable differences in lifespan between individuals in the wild and
those in captivity. In natural environments, their lifespans are generally shorter,
averaging around 10 to 15 years, due to factors such as predation, disease, and habitat
challenges. The harsh realities of the wild, including competition for food and exposure
to parasites, significantly impact their longevity. Conversely, in captivity, where
these snakes are shielded from predators and benefit from controlled diets, shelter,
and healthcare, they can live considerably longer, often reaching 20 to 30 years.
This extended lifespan in captivity is attributed to the elimination of external threats
and the provision of optimal living conditions, highlighting the impact of environmental
stressors and care on the longevity of
Lampropeltis
species.
Behavior
Lampropeltis
species are solitary and motile, displaying a range of behaviors that highlight their
adaptability and survival strategies across diverse habitats. They engage in varied
forms of locomotion, including burrowing, climbing, and swimming, to navigate their
environments effectively. Activity patterns among these snakes vary, with some being
diurnal to capitalize on cooler climates, while others adopt nocturnal habits in warmer
regions to avoid the heat. Their social interactions are mainly limited to the breeding
season, characterized by chemical communication for mating and non-lethal combat between
males for access to females. As constrictors, they employ a predation strategy that
involves suffocating their prey, allowing them to feed on a varied diet that includes
rodents, birds, and even venomous snakes, thanks to their immunity to certain venoms.
Defensive tactics include mimicry of venomous species to deter predators, showcasing
their complex interaction with the ecosystem. Overall, the behavior of
Lampropeltis
species encompasses a blend of solitary living, efficient movement, and strategic
survival techniques suited to their ecological niches.
Communication and Perception
Lampropeltis
species utilize a complex array of sensory adaptations for communication and environmental
perception, heavily relying on chemical signals and tactile interactions. Pheromones
play a pivotal role in mating, with females emitting them to attract males, who detect
these cues using the Jacobson's organ, a key component in chemical communication and
territorial awareness. Physical contact, especially during the breeding season, serves
as a form of tactile communication, where males engage in combat to establish dominance
and mating rights. Although their vision is not highly developed,
Lampropeltis
snakes compensate with sensitive vibration detection through their jawbones. Tongue
flicking is a distinctive behavior for sampling environmental chemical cues, crucial
for navigating, hunting, and avoiding predators. Together, these sensory and communication
methods underline the adaptability of
Lampropeltis
species, enabling them to thrive in diverse environments by efficiently locating
mates, identifying prey, and sensing threats. Mimicry in
Lampropeltis
species is a fascinating example of evolutionary adaptation for survival. Many
Lampropeltis
exhibit Batesian mimicry, where they mimic the coloration and patterning of venomous
snakes, such as coral snakes, despite being non-venomous themselves. This mimicry
serves as a defensive strategy, deterring predators who mistake them for the more
dangerous species they resemble. The strikingly similar color bands found in some
Lampropeltis
species and venomous counterparts effectively confuse potential threats, providing
an added layer of protection in their natural habitats. This form of mimicry not only
highlights the complex interactions within ecosystems but also underscores the adaptive
strategies developed by prey species to avoid predation.
- Other Communication Modes
- mimicry
- Perception Channels
- visual
- tactile
- acoustic
- vibrations
- chemical
Food Habits
Lampropeltis
species are carnivores with a diet that encompasses rodents, birds, eggs, other reptiles
(including venomous snakes), and amphibians, showcasing a wide prey spectrum facilitated
by specialized feeding adaptations. Their hinged jaws and expandable esophagus allow
them to consume prey significantly larger than their head, while their method of constriction
efficiently subdues and kills prey before ingestion. Notably, certain
Lampropeltis
species exhibit an immunity to venom, enabling them to prey on venomous snakes such
as rattlesnakes and copperheads without harm. This combination of dietary diversity
and physiological adaptations highlights
Lampropeltis
as versatile predators within their ecosystems, capable of exploiting a range of
food sources through unique hunting and feeding strategies.
- Primary Diet
-
carnivore
- eats terrestrial vertebrates
Predation
Lampropeltis
species exhibit a sophisticated array of anti-predator adaptations that enhance their
survival across diverse habitats. Behavioral strategies such as hiding and defensive
posturing, combined with the ability to quickly flee from threats, play a crucial
role in their defense. Physically, their muscular bodies not only aid in constriction
of prey but also in defensive struggles. Color patterns are particularly significant,
with Batesian mimicry allowing non-venomous species to imitate the appearance of venomous
snakes, deterring potential predators through deception. Cryptic coloration further
helps them blend into their surroundings, minimizing detection by predators such as
birds of prey, mammals, and larger snakes. Juvenile
Lampropeltis
rely more on camouflage and rapid growth to escape predation, gradually incorporating
more diverse strategies like mimicry and defensive displays as they mature. These
combined adaptations—behavioral, physical, and visual—form a comprehensive defense
mechanism that mitigates the risk of predation and underscores the evolutionary success
of
Lampropeltis
within their ecological niches. They have also been know to eat other snakes in
Lampropeltis
.
- Anti-predator Adaptations
- mimic
- cryptic
Ecosystem Roles
Lampropeltis
species play crucial roles in their ecosystems, serving as both predators and prey
within their ecological communities. As carnivorous predators, they help regulate
the populations of a diverse array of organisms, including rodents, birds, reptiles,
and amphibians, thereby maintaining ecological balance and preventing the overpopulation
of these species. This predatory behavior is particularly vital in controlling rodent
populations, indirectly benefiting human agricultural interests by reducing crop damage
and disease spread. Additionally, by serving as prey for larger predators such as
birds of prey, mammals, and other snakes,
Lampropeltis
contribute to the energy flow and biodiversity within their ecosystems. Their presence
and interactions within the food web underscore their importance in sustaining the
health and resilience of their environments, supporting a stable ecosystem structure
and function. While direct mutualistic or commensal relationships are not well-documented,
their indirect benefits to ecosystem health and human interests highlight their integral
role in ecological dynamics.
Economic Importance for Humans: Positive
Species in the genus
Lampropeltis
hold a nuanced yet significant relationship with human interests, primarily through
their ecological roles rather than direct economic contributions like food or shelter.
These snakes captivate scientists with their unique venom resistance and predation
patterns, offering insights into potential medical applications and ecological dynamics.
Ecotourism benefits from their presence, as their vibrant patterns and behaviors attract
wildlife enthusiasts, contributing to local economies. Agriculturally, they serve
as natural pest controllers, preying on rodents and reducing the need for chemical
pesticides, thereby supporting crop health and yield. Additionally, their popularity
in the pet trade highlights the importance of sustainable practices to prevent negative
impacts on wild populations. Beyond these direct interactions, the broader ecological
functions of
Lampropeltis
species, such as maintaining prey population balance and indicating healthy ecosystems,
underscore their indirect but vital contributions to human economies and environmental
health. Through these roles,
Lampropeltis
exemplify the complex ways in which wildlife can influence human economic and ecological
landscapes, emphasizing the importance of their conservation and sustainable management.
- Positive Impacts
- pet trade
- ecotourism
- research and education
- controls pest population
Economic Importance for Humans: Negative
There are no known adverse effects of
Lampropeltis
species on humans
Conservation Status
Lampropeltis
species have varied conservation statuses, ranging from Least Concern to more vulnerable
categories due to habitat loss, road mortality, and the pet trade. For instance,
Lampropeltis getula
and
Lampropeltis triangulum
are considered of Least Concern but face threats like habitat destruction and misidentification
killings. Meanwhile,
Lampropeltis zonata
is under conservation concern in specific locales due to habitat loss and illegal
collection. Conservation actions across the genus aim at habitat preservation, legal
protections, roadkill mitigation, and public education to support these species. Efforts
such as habitat restoration, implementing legal measures to limit collection, constructing
wildlife crossings, and raising public awareness about the ecological importance of
these snakes are crucial for their sustained survival and ecological balance.
Other Comments
Lampropeltis
species are characterized by their vibrant patterns and significant ecological roles.
They are steeped in scientific, cultural, and historical significance. Derived from
Greek words meaning "shiny shield," their name reflects the snakes' distinctive, radiant
scales. While the fossil record of
Lampropeltis
is sparse due to the delicate nature of snake skeletons, these snakes are part of
the colubrid family's broad evolutionary narrative, dating back to the late Eocene.
Beyond biology, species within this genus have permeated human culture and mythology,
symbolizing various concepts from healing to fear across different societies. Scientific
research into
Lampropeltis
offers insights into their adaptability, including venom resistance and coloration
patterns, crucial for understanding their survival strategies and biodiversity.
Additional Links
Contributors
Kayvon Mostofi (author), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Neotropical
-
living in the southern part of the New World. In other words, Central and South America.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- desert or dunes
-
in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- chaparral
-
Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- rainforest
-
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
- scrub forest
-
scrub forests develop in areas that experience dry seasons.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- urban
-
living in cities and large towns, landscapes dominated by human structures and activity.
- suburban
-
living in residential areas on the outskirts of large cities or towns.
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- indeterminate growth
-
Animals with indeterminate growth continue to grow throughout their lives.
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- year-round breeding
-
breeding takes place throughout the year
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- internal fertilization
-
fertilization takes place within the female's body
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- young precocial
-
young are relatively well-developed when born
- diurnal
-
- active during the day, 2. lasting for one day.
- nocturnal
-
active during the night
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- hibernation
-
the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.
- solitary
-
lives alone
- visual
-
uses sight to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- mimicry
-
imitates a communication signal or appearance of another kind of organism
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- vibrations
-
movements of a hard surface that are produced by animals as signals to others
- chemical
-
uses smells or other chemicals to communicate
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- pet trade
-
the business of buying and selling animals for people to keep in their homes as pets.
- ecotourism
-
humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.
- carnivore
-
an animal that mainly eats meat
References
Blanchard, F. 1921. A revision of the king snakes genus Lampropeltis . United States National Museum: Smithsonian Institution.
Bryson Jr., R., J. Pastorini, F. Burbrink, M. Forstner. 2007. A phylogeny of the Lampropeltis mexicana complex (Serpentes: Colubridae) based on mitochondrial DNA sequences suggests evidence for species-level polyphyly within Lampropeltis. Molecular Phylogenetics and Evolution , 43(2): 674-684.
Burbrink, F., H. Yao, M. Ingrasci, R. Bryson Jr., T. Guiher, S. Ruane. 2011. Speciation at the Mogollon Rim in the Arizona Mountain Kingsnake (Lampropeltis pyromelana). Molecular Phylogenetics and Evolution , 60(3): 445-454.
Dessaue, H., F. Pough. 1975. Geographic variation of blood proteins and the systematics of kingsnakes (Lampropeltis getulus). Comparative Biochemistry and Physiology Part B: Comparative Biochemistry , 50(1): 9-10.
Ernst, C., E. Ernst. 2003. Snakes of the United States and Canada . Smithsonian Books: Smithsonian Books.
Judd, W., K. Krysko. 2006. Morphological systematics of kingsnakes, Lampropeltis getula complex (Serpentes: Colubridae), in the eastern United States. Zootaxa , 1193: 1–39.
Kohler, D., B. Hamilton, D. Dittmer, A. Whiting. 2023. Citizen Science in Action: An Updated Distribution for Lampropeltis pyromelana. Western North American naturalist , 83(2): 165-175.
Myers, E., J. RodrĂguez-Robles, D. DeNardo, R. Staub, A. Stropoli, S. Ruane, F. Burbrink. 2013. Multilocus phylogeographic assessment of the California Mountain Kingsnake (Lampropeltis zonata) suggests alternative patterns of diversification for the California Floristic Province. Molecular Ecology , 22(21): 5418-5429.
Penning, D., B. Moon. 2017. The king of snakes: performance and morphology of intraguild predators ( Lampropeltis ) and their prey ( Pantherophis ). Journal of experimental biology , 220(6): 1154–1161.
RodrĂguez-Robles, J., D. DeNardo, R. Staub. 1999. Phylogeography of the California mountain kingsnake, Lampropeltis zonata (Colubridae). Molecular Ecology , 8(11): 1923-1934.
Ruane, S., . Bryson Jr., R. Pyron, F. Burbrink. 2014. Coalescent Species Delimitation in Milksnakes (Genus Lampropeltis) and Impacts on Phylogenetic Comparative Analyses. Systematic biology , 63(2): 231-250.
Ruane, S. 2015. Using geometric morphometrics for integrative taxonomy: an examination of head shapes of milksnakes (genus Lampropeltis ): Geometric Morphometrics of Milksnakes. Zoological journal of the Linnean Society , 174(2): 394-413.